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HARVARD UNIVERSITY
e
Library of the
Museum of
Comparative Zoology
The Systematics of Neotropical
Orb-weaving Spiders in the
Genus Metepeira (Araneae: Araneidae
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. PIEL'
CONTENTS
FANS trac tees See LEER reed PA ae LY, 1
Imtrocductionie 2... 2e5 ao ae ek ee 2,
Acknowledgments) #2 esses en ei ee neat ae 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 stoplemale Metepeind, yo sss. se seen ee 12
Key ston MialesMetenet ia te saws. se ee 2 lef
IMICLenetnauoxts Group les saree mele. ewes Is)
1. Metepeira datona Chamberlin and Ivie 20
2. Metepeira desenderi Baert ___........ ail
3. Metepeira grandiosa grandiosa
Chamberlin and Ivie 23
4. Metepeira grandiosa alpina
Chamberlin and Ivie _ 24
Metenevrawicilasc: Group ese elo cian 26
5. Metepeira cajabamba New Species _.. 26
6. Metepeira glomerabilis (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 Oil
Metepeira nigriventris Group —.........--.--------------- 38
11. Metepeira nigriventris (Taczanowski) 38
12. Metepeira tarapaca New Species _..... 40
13. Metepeira calamuchita New Species. 42
14. Metepeira galatheae (Thorell) 43
15. Metepeira karkii (Tullgren) 46
iMeteneinascompsa Croup =: at en eee 47
16. Metepeira compsa (Chamberlin) —__ 48
17. Metepeira roraima New Species —-...... D3
18. Metepeira gressa (Keyserling) — 54
Metepeira incrassata Group —.2--- 56
19. Metepeira maya New Species __.......... 56
20. Metepeira inca New Species -............... 58
‘Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts 02138. Current
address: Institute of Evolutionary and Ecological Sci-
ences, Leiden University, 2311 GP Leiden, The
Netherlands; piel@rulsfb.leidenuniv.nl.
21. Metepeira gosoga Chamberlin and Ivie
Atcha Cadre Some iligteeetye tale SON ceed Siete teeta Te 59
22. Metepeira olmec New SESS 60.
23. Metepeira comanche Levi ____------- 62
24. Metepeira pimungan New Species ..... 62
25. Metepeira triangularis (Franganillo) _ 63
26. Metepeira arizonica Chamberlin and
[vie tir ee aie Peasy sehen as 6 Ahead Bry) 66
27. Metepeira atascadero New Species ~.. 67
28. Metepeira incrassata F. O. P.-
Cambridge yrs ene. pales ee ee 68
Metepeira ventura Group 20.0222 71
29. Metepeira ventura Chamberlin and
vi See Renee el ET ae eee re: ee Bk 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
[hvalevnael Scie oe RM Den OLB ayiee elem ccd RELY a
34. Metepeira chilapae Chamberlin and
vac jos epee ae Wee ae ee eee ee 78
Metepeira minima Group ~....--..--.------------------------- 80
35. Metepeira petatlan New Species —...... 80
36. Metepeira minima Gertsch _............-.- 82,
37. Metepeira pacifica New Species 84
38. Metepeira jamaicensis Archer 86
[Eaten abinen (Cue) 88
Ds Gl x es ee sae RE ree | Eu 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. cajabamba, M. calamuchi-
ta, M. celestun, M. inca, M. lacandon, M. maya, M.
olmec, 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 il
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. triangularis, M. chilapae,
M. comanche, M. compsa, M. crassipes, M. datona,
M. desenderi, M. galatheae, M. glomerabilis, 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 labyrin-
thea. Levis (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
This paper is part of my Ph.D. thesis for
the Department of Organismic and Evo-
lutionary Biology, Harvard University. Iam
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
Fitzpatrick, Germania Jacome, Ant6nia
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 every 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
American Museum of Natural
History, New York, United
States; N. Platnick, L. Sorkin
Natural History Museum, Lon-
don, England; P. Hillyard
California Academy of Sciences,
San Francisco, California, Unit-
ed States; C. Griswold
Carlos Valderrama A.; Bogota,
Colombia
Florida State Collection of Ar-
thropods, Gainesville, Florida,
United States; G. B. Edwards
Institut Royal des Sciences Na-
AMNH
BMNH
CAS
CV
FSCA
IRSNB
JAK
JEC
JMM
MACN
MCN
MCZ
MECN
MEG
MLJC
MLP
MNRJ
MNSD
MUSM
MZSP
MZUF
NRMS
PAN
METEPEIRA ° Piel 3
turelles de Belgique, Brussels,
Belgium; L. Baert
J. A. Kochalka, Ciudad Univer-
sitaria, Paraguay
J. Carico, Lynchburg, Virginia,
United States
J. Maes, Le6n, Nicaragua
Museo Argentino de Ciencias
Naturales, Buenos Aires, Argen-
tina; E. A. Maury, C. L. Scioscia
Museu de Ciéncias Naturais,
Fundagao Zoobotanica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil; E. H.
Buckup, M. A. L. Marques
Museum of Comparative Zool-
ogy, Harvard University, Cam-
bridge, Massachusetts, United
States: H. W. Levi
Museo Ecuatoriano de Ciencias
Naturales, Quito, Ecuador; Ger-
mania Eistévez Jacome
M. E. Galiano, Buenos Aires,
Argentina
Maria Luisa Jiménez, Centro de
Investigaciones Bioldgicas del
Noroeste, La Paz, Mexico
Museo de Universidad Nacional,
La Plata, Argentina; R. F. Arro-
zpide, C. Sutton
Museu Nacional, Rio de Janeiro,
Brazil; A. Timotheo da Costa
Museo Nacional de Historia
Natural, Santo Domingo, Re-
publica Dominicana; Félix Del
Monte
Museo de Historia Natural,
Universidad Nacional Mayor de
San Marcos, Lima, Peru; D. Silva
Museu de Zoologia, Universida-
de de Sio Paulo, Sao Paulo, SP,
Brazil; P. Vanzolini, J. L. Leme
Museo Zoologico de “La Spe-
cola” Universita di Firenze,
Florence, Italy; S. Whitman
Naturhistoriska Riksmuseet,
Stockholm, Sweden; T. Krones-
tedt
Polska Akademia Nauk, Warsza-
4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
wa, Poland; J. Prészynski, A. Slo-
jewska, W. B. Jedryczkowski
R. E. Leech, Edmonton, Alber-
ta, Canada
Forschungsinstitut Sencken-
berg, Frankfurt am Main, Ger-
many; M. Grasshoff
Susan Riechert, Knoxville, Ten-
nessee, United States
National Museum of Natural
History, Smithsonian Institution,
Washington, D.C., United
States; J. Coddington, S. F.
Larcher
Zoologisches Museum der
Humboldt Universitat, Berlin,
Germany; M. Moritz
Zoologisk Museum, Copenha-
gen, Denmark; H. Enghoff, N.
Scharff
Zoologische Staatssammlung,
Munich, Germany
REL
SMF
SR
USNM
ZMB
ZMUC
ZSM
Locality Data Storage and Manipula-
tion. Locality data from each collection vial
were entered into a database designed us-
ing Claris 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/ens/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
made 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 DMAAC ONC
aeronautical maps; in most cases, elevation
was estimated using NOAA data with an
on-line database server (http://phylogeny.
harvard.edu/~piel/find. html).
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 activi
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 sympatric 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 CCD
video camera, chosen for its high sensitiv-
ity to light. Video images were captured
using a QuickImage™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 Xanté™ 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 sclerites. 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 OmniChrom™ pencil and a
drafting pen with India ink and then was
scanned at 600 dpi on a LaCie Silverscan-
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
METEPEIRA ° Piel 5
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
web forms scaffolding around an almost
vertical orb and supports the spider's 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 lighter re-
gion takes up almost 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 pigmented 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 sternum. 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 show hints of white markings
in some specimens. Characteristics found
in the carapace, abdomen, and sternum of
Metepeira are also found in Araneus
koepckeorum 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).
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, Zygiella, 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 ri
Barrier Web White Dorsal Light Eye Region
/ Fleur-de-Lis
Pattern
Median White
Line on Sternum
See \\\ , * if _ <
We HUAI ROT SES Sa \ Median White
it BS SVANS TS 7 D 3 Line on Black
Venter
Orb Web
nigriventris incrassata minima
labyrinthea compsa ventura
Eee
" —— — Flagellae
Dy Lae \Y7 ‘on Stalk (+)
: DEA (-)
K on MA (-)
F, Teeth on Face of K (+)
DEA (+) | North America
“
SC fEnlarged TA (+) :
; [) South America
datona
(1)
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 labyrinthea
(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; TD, 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 the anterior side of the
femur, and between zero and seven on the
anteroventral side. Males typically have
more 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 male 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, Metepeira’s 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 terminal 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). Some-
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 the 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 Cyrtophora 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 of Cyrtophora 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-
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. pimungan 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 communication), M. tarapaca (VN.
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-
METEPEIRA ° Piel 9
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);
dry 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. compsa) 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. 1
from the equator (e.g., M. rectangula, M.
vigilax, M. cajabamba, Fig. 36; M. arizon-
CG, See 2i3)):
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 Mecynogea
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. celestun
(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. glomerabilis and M. vigilax;
M. compsa and M. glomerabilis; M. comp-
sa and M. nigriventris; M. conypsa and M.
inca; 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.
bensoni. Jiménez 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 lindae, 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 incompatible
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. cajabamba,
M. glomerabilis, M. rectangula) 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
“al
e datona S
= desenderi
Map|
Map 3
METEPEIRBA ° Piel 11
\
= grandiosa alpina ¢
e grandiosa grandiosa ~~
Se
@ vigilax
| + rectangula
/
\e ns j My
f Se
Map 4 i 2
Maps 1, 2. Metepeira foxi 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
1 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) pe ee IE EY eR eck. \od OL ONE earn 9}
= 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)
Epigynal openings shaped like the en-
trance to a snail shell (Fig. 55); His-
paniola, Brazil, Argentina (Map 4)
Be ER ee eee ee (7) vigilax
= Epigynal openings tubular (Figs. 40, 48)
Epigynal openings strongly tilted (Fig.
40) Pexul(Mapro)s = a (5) cajabamba
= Epigynal openings weakly tilted (Fig. 48);
Colombia to Brazil (Map 3) ———---
Sia EL We SSN Smee nee dat REO (6) glomerabilis
Epigynal openings large and gaping, cre-
ating large atria inside (Figs. 61, 62);
Chile and Argentina (Map 4) ————
Seer Se eS ee (8) rectangula
= Epigynal openings not gaping and not
creating large atria (e.g., Fig. 69) ______.. 5
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 (eseeehigs ss lOlen 42) sa eeeenes 11
Sternum with longitudinal white line
(Fig. 21); Galapagos Islands (Map 1)
yall ak ISN ae Aen al (2) desenderi
~ Sternum entirely black or brown (Figs.
Igy OAS wis)
Abdomen wider than long (Fig. 12); Flor-
ida to Hispaniola (Map 1) _- (1) datona
= Abdomen longer than wide (Figs. 27, 34)
Coxae as black as sternum (Levi, 1977,
fig. 98); Canada—U.S. border (Levi,
ESO Ch suey o\\))) akin ee
Delia (Levi, 1977: 212) grandiosa palustris
= Coxae yellow or orange and lighter than
Stemmum(Higsy28.5.3.0) a) as eee 9
Epigynum with crescent-shaped sclero-
tized openings on either side of the
scape (Levi, 1977, fig. 87); western
U.S. and Canada (Levi, 1977, map 2)
Pesce ses Ayr Os (Levi, 1977: 210) foxi
= Epigynum with wide, transverse depres-
sion (Figs. 23, 24, 30, 31)
Scape wide and stubby (Fig. 24); Baja
California north to Canada (Map 2) —
BSI 5 Shad 2 (3) grandiosa grandiosa
— Scape triangular (Fig. 31); in mountains
from north-central Mexico to Canada
(Mapa?) eee aes (4) grandiosa alpina
Scape thickness equal to greater than
width of epigynal depressions (e.g.,
Figs. 86, 119, 131, 143, 180, 201, 266)
9(8)
10(9)
11(5)
= Scape narrower than epigynal depres-
sions, or epigynal depressions in the
shape of longitudinal slits (e.g., Figs.
D3 8¥ 2520230) iain cies ks Se eae ana 35
12(11) Base of scape originates anteriorly and
projects ventrally before curving 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)
= Scape does not create a noticeable gap or
overhang (e.g., Figs. 77, 78, 130, 131,
NGS GGMO6 SIN Veit te eine ee eens 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.
OD )\eees 3 = aR A a ee 14
14(13) Sternum black, coxae mostly black (Fig.
A calamuchita
g@ karkii
@ nigriventris
# compsa
® roraima
@ gressa
mi Skangse
A PN as, Rg
METEPEIBA ° Piel 13
Maps 5, 7. Metepeira nigriventris species group. M. calamuchita, M. karkii, M. nigriventris, M. galatheae, M. tarapaca.
Map 6. Metepeira labyrinthea species group. M. spinipes, M. lacandon.
Map 8. Metepeira compsa species group. M. compsa, M. roraima, M. gressa.
91), and carapace without lighter me- 15(14) Stermum brown to black with parallel
dian mark (Fig. 90); high altitudes in
Bolivia and Peru near Lake Titicaca
(IMfayp 35) hy Wetec ee AE aol ae (11) nigriventris
= 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)
lines on either side of median white
line on venter: the parallel lines are
thicker 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,
AUMIES Seite? (0) PRs. CA eee oS, ee 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 wrinkled portion of the
scape’s 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-
tural (ESI 9.539 yp) 2) eee ae eee ee ee 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
(MIEN oe) ieee a (13) calamuchita
— General shape of epigynum square.
Scooped-out depressions project more
laterally than posteriorly (Fig. 94);
northern Chile and southern Peru
(Miapiii)) ea Sot ke Se ee (12) tarapaca
18(12) Epigynal openings almond-shaped, not
noticeably sclerotized, and at a 40° to
60° angle from axis of spider (Figs. 69,
cS) ise ene LES Saat. CE Awaba aut aie 19
= Epigynal openings not almond-shaped
(eg, Figs. 180, 187, 201), or if al-
mond-shaped, slightly sclerotized and
at an angle from spider's axis of 80° to
100° (larress, IMSL ISIS IGi6) 2)
19(18) Almond-shaped openings created by de-
pression where scape arises from epi-
gynum (Fig. 173); California, Arizona,
northwestern Mexico (Map 9)
Se a = Se ee eee a (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 the 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
GOUCS) KYOTOLE TH SIME 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
the 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
the 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) ———--——
Spe cee tah Se lsP PEs (Levi, 1977: 196) labyrinthea
= For 125 mtDNA sequence data, the fol-
lowing sequences are diagnostic. Base
14261: ACGAT;: 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
CH Rpereereeeeee CAE wi ROM mM INC n ls ns ou" (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
(Miayp.S)) 2 ia eee) Sees (16) compsa
= Epigynal openings not small, round, and
SCIETOLZE Ci, ee Wan ees nee ie eee 23
23(22) Rim of epigynal openings sclerotized and
in an oval or teardrop shape (Figs. 151,
5 OSA GGE AUST srsstaet 6 es Rien ee ee 9A
- Rim of epigynal openings not sclerotized
in an oval or teardrop shape (Figs. 180,
NOAL 9 Oilise2, 08) Ls Ai eee OAT
24(23) Epigynal openings oval, small, and partly
hidden by scape (Fig. 151); northern
Argentina, Uruguay, and southern Bra-
Zilt (Miaps8)\ see ener t re ied (18) gressa
— Epigynal openings teardrop-shaped (Fig.
187) or large and oval but not hidden
loyy Sroatoxsy (laress, ISS), TGS)
25(24) Epigynal openings teardrop-shaped (Fig.
187); northeastern Mexico to Texas
(Kevin oiieemapeleMap >) ==
ee (23) comanche
= Epigynal openings oval-shaped (Figs.
IL) OS SG. G)) sieesas. hs Cen eae ee eee SUE 26
26(25) Lower lip of epigynum pointed (Fig.
166), abdomen white (Fig. 169); north-
emmytipyorkerul (Mapa lil) james (20) inca
METEPEIRA ® Piel 15
+ arizonica ~~
@ gosoga ce
Bal ais @ pacifica
% inca
olmec
incrassata mg petatlan
@ pimungan
Bre
Map 9 Map 10 Map 11
ta
@ uncata w ventura
@ crassipes a 4 celestun
g revillagigedo ® chilapae
@ jamaicensis
B minima
# atascadero
® triangularis
Map 14
Maps 9, 11, 14. Metepeira incrassata species group. 9, M. arizonica, M. gosoga, M. incrassata, M. pimungan. 11, 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 or, if sclerotized, not in shape of sun-
pointed (Fig. 159), abdomen dark (Fig. glasses) (ce: Higs1208.( 224) ieee ee 28
162); southern Mexico and Belize to 28(27) postenion epigynal lobes converge behind
Costa Rica (Map 14) ___.... (19) maya the scape so that epigynal depressions
27(23) Rim of epigynal openings sclerotized and appear closed off from ventral view
shaped like a pair of sunglasses (Fig. (HigswilS08 208521160224) aia a 29
201); eastern Cuba and Hispaniola = Posterior epigynal lobes end before they
(Mapp rll) ees re hh. (25) triangularis disappear behind the scape so that epi-
= Rim of epigynal openings not sclerotized gynal depressions appear open poste-
16 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
riorly from ventral view (Figs. 194,
DAD. Oko 6 6) ea eran ie eed ne A a 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)
BAS Mt 60 Aine on Bere em (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. 2082268224) ee eens 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
Oe tie ee SOR. EME eel eels (26) arizonica
= Epigynum not puffy, scape not so fat that
depressions become crescent-shaped
(Bigs DIGHOOA) ea tah tine ven 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)
fete Biri tithe Serene eRe ey it's CoP (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
IAS) mrs rae eee een Re ee (27) atascadero
32(28) Small square-shaped epigynal depres-
sions on either side of scape (Figs. 259,
DGG) pte vttts eased ook kale Reisen oar Sire end? cate i)
= Large, more rounded epigynal depres-
sions (Fig. 194) or with straight to S-
shaped edges mostly covered by scape
(Bigs 245) iernot darstinvie Ni oi8 cella 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
(Mia ppl) isiat as eee ae ee (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. 1:2) fs teas a At se ties (33) crassipes
34(32) Large, rounded epigynal depressions
with large black spheres inside (Fig.
194). Sternum black with posterior
dewdrop-shaped mark. Pair 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
(Mato 1:3) ate en career (31) celestun
Epigynal depressions wider than long
(Figs, 23015059) 1b oN Wiel cobae 36
Epigynal depressions longer than wide
(e.g., Figs. 238, 273, 280, 288) 37
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
(Miao: S))ant= es ONE eee (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
(Mayon 12) 2a else besornei ty byee (32) uncata
Epigynal depressions indistinct anterior-
ly. Black comma-shaped marks inside
depressions and covered by translucent
membranes (Fig. 238); Isla Socorro of
the Archipiélago de Revillagigedo
(Mapel2)ie 2s ee (30) revillagigedo
Epigynal depressions distinct anteriorly.
Black spheres shifted laterally and lo-
cated outside the depressions (e.g.,
Figs. 273, 280, DOS) gree See eae bat ees: 38
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
Scapen(Hicsh:27/3 92.05) yaeeeeann aaeeeaennens 39
Dark spheres larger than epigynal open-
ings (Fig. 273); west coastal Mexico
(Mapytl 0) ees ree a (35) petatlan
Dark spheres smaller than epigynal open-
Ingsa(MICSHZO S29 5) pees een ernene 40
V-shaped ridge under the 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) ——
se te et See Pe (37) pacifica
Straight ridge under the scape. Dark
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
Terminal apophysis thin, small, fleshy,
without teeth or sclerotized parts (Figs.
OES ES SOR OAS)) | kara Rai tae Lae Meee. 5 RE 2,
Terminal apophysis enlarged, meaty, with
teeth or sclerotized parts (e.g., Figs.
3 SMO ORSAT a9 9) eee ee ee 7
Terminal apophysis narrow; embolus
curled clockwise like the tip on a cork-
screw (Figs. 7, 15)
Terminal apophysis wide; embolus tilted
up and L-shaped (Figs. 22, 29) 4
Curled embolus with a ridge on the up-
per surface and raised on a pedicel;
basal embolic apophysis enormous
(Fig. 15). Sternum with wide median
white line (Fig. 21); Galapagos Islands
(AWE T opal) Sree eee een es (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)
SS ha ae iene NT (1) datona
L-shaped embolus at an acute (< 90°) an-
gle (Levi, 1977, figs. 91-93); western
U.S. and Canada (Levi, 1977, map 2)
Be ies robes i (Levi, 1977: 210) foxi
L-shaped embolus at angle of 90° or
preate;rs (igs# 22-229) Wate Se ees 5
Median apophysis with rounded projec-
tion on dorsal side; jagged posterior
edeejoiikeel (Bigsk 220029), sas 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)
eid es (Levi, 1977: 212) grandiosa palustris
Lower, transverse part of L-shaped em-
bolus longer than vertical part (Fig.
29): mountains from north-central
Mexico to Canada (Map 2) —-—---—
pee De Sets Sees See ene (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
Terminal apophysis does not entirely
overhang the embolus. Embolus long
and robust, with a long and thin gap
created between the embolus and the
11(7)
14(13)
15(12)
METEPEIRA ° Piel 17
basal embolic apophysis (Figs. 38, 46,
53, 60)
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)
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
(HiGS38. 465.60) eee dt clio 9
Keel absent or greatly reduced (Fig. 38);
Peru. (Map 3) ae (5) cajabamba
Keel present (Figs. 46, 60)
Large embolus as long as basal embolic
apophysis (Fig. 60). Normal embolic
cap; Chile and Argentina (Map 4) ——
es ee ieee RENE aSAL) (8) rectangula
Small embolus shorter than basal embol-
ic apophysis, often seen with winged
embolic cap (Fig. 46); Colombia to
Brazile(Map! 3) 2. ses (6) glomerabilis
Median apophysis with keel (e.g., Figs.
QONIAT py SEs cele Rok aan ee Seer ee 12
Median apophysis without keel (e.g.,
Figs. 157, 178, 286); all North Ameri-
can or Caribbean species except for M.
UVC CAEN SRS et eS 29;
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
Distal embolic apophysis sleek, pointed,
and feather-shaped when viewed from
underside of terminal division (Fig.
70); Mexico City north to California
(OMEN op 6) ak te eee ees (9) spinipes
Distal embolic apophysis spoon-shaped
(Fig. 6) or widened with bump (Fig.
79)
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) labyrinthea
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
Distal embolic apophysis a simple exten-
sion that projects forward, parallel to
the embolus. Keel usually rounded
18
16(15)
17(16)
18(17)
19(18)
20(19)
21(16)
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,
TOO, WSLS WAL TANS) ee 16
Distal embolic apophysis points up and
away from the embolus. Keel usually
jagceda(Euigsael 4k 4Q) ieee eee 21
Distal embolic apophysis wide and raised
up on boss. Keel usually pointed (Figs.
SARAO DSi OO) eater oad eh a eee 17
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)
ESSE RRO Soe i Tes (13) calamuchita
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,
Ot Meili QA) Meant telat en dete cle
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.
Sie) Spell ee loa ta aE te
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
(ESS pS i159.) ee aie eee Berchet rah hele
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)
osc enone ena (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
Distal embolic apophysis thinner than
embolus near the junction of the two.
Embolus tip is curved gently (Fig.
20
ES
bo
ey)
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
Distal embolic apophysis a projecting
bumpy (hig ste oi 64 5oli/fll) eeneeern na 23
Distal embolic apophysis curved off
(Figs. 178, 185, 199, 214), rounded
(Figs. 192, 206), or absent (Figs. 264,
DASE 29 Bhs. NEA Rates. Sel eae teat AE
Distal embolic apophysis raised up from
the embolus and pointed (Fig. 171);
California, Arizona, northwestern Mex-
Tato) (NYY oy 0) a ee (21) gosoga
Distal embolic apophysis not raised up,
but projecting forward and rounded off
(Bigs.057 164). 0 a Wa
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) maya
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-
25
24
teaua(Mapel4) aa (27) atascadero
Embolus without such bump near tip
(Ces AhIE R222) AGS eee ee 26
Distal embolic apophysis abruptly ends
in sharp curve and flagellae not set off
on a narrow stalk (Figs. 178, 185, 199,
DOO Ni tht S rule Sean tibet ee SOURS eaters een see ik
Distal embolic apophysis gently curved
off (Figs. 229, 236, 243), rounded
(Figs. 192, 206), or absent (e.g., Fig.
DO BN) (52! wae ake betel erred aa ee
Flagellae very 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)
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.
178, 185). Sternum with median white
Markie sa (IGS gl S25 GQ) asec e ns 29
= Highly social species. Height of embolus
plus distal embolic apophysis just be-
fore the latter curves off sharply 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)
oP NOS eee (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,
lOv@eemape "Mapa ie es (23) comanche
30(26) Distal embolic apophysis rounded off to
form convex curve (Figs. 192, 206) _..... 31
= Distal embolic apophysis gently falls off
to form concave shape (Figs. 929, 236,
243)! or absent (e:¢., Fig.) 293) i222 32
31(30) Embolus S-shaped (Fig. 192). Sternum
black with posterior white mark. Ven-
ter without white anchor shape mark
(Fig. 196); San Nicolas Island off
southern California (Map 9) —-—-———
See eee ee a ES (24) pimungan
= 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,
ORNG Nee eae ee Sa ae eee 38
~ Flagellae on median apophysis not set off
on a separate, narrow, stalk (e.g., Figs.
99959 GA) et 2 eRe OU) aa eer 33
33(32) Larger flagellum twice as wide as smaller
flagellum (Fig. 236); Isla Socorro of the
Archipiélago de Revillagigedo (Map
UT: ip ater esate Bene fe ttt sede a (30) revillagigedo
= Larger flagellum less than twice as thick
as smaller flagellum (e.g., Figs. 229,
GALEN Re Tienes Beter Mie Laut, aan eee 34
34(33) Flagellae on median apophysis relatively
(dawn) (URakas PABKO), PAGV)) Le ee 35
- Flagellae on median apophysis thicker
(Figs. 2.2 QUOTAS UES 517) es seemed Me ake EERE 36
35(34) Embolus with sharp bend near the tip
METEPEIRBA ° Piel 19
(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 mostly
black with small white mark at poste-
rior end of sternum (Fig. 254); Gua-
temala and Costa Rica (Map 12) ———
DDS 5 RE (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
thestipa(Hics 32430 2517) mass ears wees 37
37(36) Sclerotized portion of embolus about as
long as the longer flagellum (Fig. 243);
Yucatan Peninsula (Map 13) ————
tee ata Rot Die 2 oa Salen Kom SB (31) celestun
= Sclerotized portion of embolus shorter
than the longer flagellum (Fig. 257);
northwestern Mexico and California
(Miers i) ea eee ae Rea (33) crassipes
38(32) Embolus thick and tapering to a point
(Figs. SATA OAT S)) |peecmeretias Wit S MeN ESE hy pee ROE 39
= Embolus thin and needlelike (Figs. 286,
DAC HN a heee tee eee en Moen nee Se ea Oe, 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 ells) ies: sere pe Stee (36) minima
40(38) Flagellae set off on long, thin stalk (Fig.
293); Cayman Islands, Jamaica, and
laity (Miappilis)) a seee aes (38) jamaicensis
= Flagellae set off on short, thicker stalk
(Fig. 286); Honduras to Costa Rica
(Miay sli) gcse Be. A ewes ote 8 (37) pacifica
Metepeira foxi Group
The M. foxi species group (sensu Levi,
1977) and M. desenderi share very similar
genitalia, 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 Zoology, 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.
OF 7. 2473 witha Figs. 143, 252),
1. Metepeira datona
Chamberlin and Ivie
Figures 7-13, 312; Map 1
Metepeira datona Chamberlin and Ivie, 1942: 68, fig.
196, 2. Female holotype from Daytona Beach,
Florida, USA, in the AMNH, examined. Levi,
1977: 208-210, figs. [S=8 OO Rn Brignoli, 1983:
Dp Ds
Metepeira inerma Bryant, 1945: 378. Female holo-
type from Cap Haitien, Haiti in the MCZ, exam-
ined. First synonymized by Levi, 1977: 208-210.
Description. Female from Morant
Point, Mammee Bay, Jamaica. Carapace
light aroma 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 sometimes 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 an tibia 2.4,
metatarsus 1.5, tarsus 0.8. Second patella
and tibia 1.9 mm, 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 female’s 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 curves 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)
buat unlike M. grandiosa grandiosa (Fig.
22) and M. grandiosa alpina (Fig. 29), the
posteroventral edge of the keel on the me-
dian apophysis is rounded, as opposed to
pointed (Fig. 7). Finally, M. datona has the
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
i)
Records Examined. BAHAMAS Abaco Cays: Aba-
co, 26°29'N, 77°5'W, 2.xii.1964 (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, 926°43'N, 77°14'W,
121.1964 (W. B. Peck, CAS). Acklin’s Id.: Atwood’s
Harbor, 22°13’N, 74°18'W, 15.ix.1958 (A. W. Scott Jr.,
MCZ): Salina Point, 22°13’N, 74°18’W, 15.viii.1958
(R. Robertson & A. W. Scott Jr, MCZ). Andros: Fresh
Creek, 24°26'N, 77°57’W, 23.iv.1953 (L. Giovannoli,
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.1967 (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, 1.v.1953 (Hayden & Giovannoli,
AMNH). Crooked Island: North shore of Cripple
Hill, 22°49'N, 74°16’W, 15.ix.1958 (A. W. Scott Jr,
MCZ): NW end, Gordon (= Gun) Bluff, 22°50'N,
74°20'W, 15.viii.1958 (R. Robinson & A. W. Scott Jr,
MCZ). Crooked Island Group: Long Cay, 22°37'N,
74°20'W, 7.iii.1953 (Hayden & Giovannoli, AMNH).
East Plana Cay: 22°37'N, 73°33'W, 4.iii.1953 (E.
Hayden, AMNH). Exumas: Musha Cay, 23°50'N,
76°15'W, 29.xii.1985 (A. Boutard, MCZ): Warderick
Wells Cay, 24°22'N, 76°36'W, 9.i1.1953 (L. Giovannoli,
AMNH), 11.i.1953 (Hayden & Giovannoli, 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 [P],
25.vii.1965 (L. Pinter, MCZ). Long Island: Clarence
Town, 23°6'N, 74°59'W, 10.iii.1953 (L. Giovannoli,
AMNH); Deadman’s Cay, 23°14’N, 75°14'W,
11.iii1.1953 (E. Hayden, AMNH). New Providence Is-
land: 7 mi W. of Nassau, 25°5’N, 77°28’W, 4.1.1953
(Hayden & Giovannoli, AMNH). North Bimini:
25°44'N, 79°15'W, 25.i.1950 (C. M. Bogert, AMNB#),
METEPEIRA ° Piel 21
6.vi. 1950 (M. Cazier & F. Rindge, AMNH), 15.v.1951
(W. J. Gertsch & M. Cazier, AMNH), 15.vi.1951 (M.
Cazier, P. & C. Vaurie, AMNH), 15.vii.1951 (C. & P.
Vaurie, AMNH), 13.xii.1952 (A. M. Nadler, AMNH),
28.xi.1959 (A. M. Nadler, AMNH). Rum Cay: near
Port Nelson, 23°38'N, 74°50'W, 16.iii.1953 (Hayden
& Giovannoli, AMNH). South Bimini: 25°42'N,
7T9°17'W, 12.vi.1950 (M. Cazier & F. Rindge,
AMNH), 22.vi.1950 (M. Cazier & F. Rindge,
AMNH), 15.v.1951 (W. J. Gertsch & M. Cazier,
AMNHB), 15.vii.1951 (C. & P. Vaurie, AMNH),
A viii.1951 (C. & P. Vaurie, AMNH), 4.xii.1952 (A. M.
Nadler, AMNH), 25.iii.1953 (A. M. Nadler, AMNH);
Gun Cay, 95°35'N, 79°20'W, 15.vi.1951 (AMNH).
BRITISH WEST INDIES Caicos Islands: Long Cay,
21°28'N, 71°33'W, 10.ii.1953 (E. Hayden, AMNH);:
South Caicos, from webs in upper beach zone,
21°31'N, 71°30'W, 3.iv.1973 (D. W. Buden, MCZ);
West Caicos, 21°39'N, 72°28’W, 4.ii.1953 (Hayden &
Giovannoli, AMNH), 5.ii.1953 (Hayden, Rabb, &
Giovannoli, AMNH). Grand Cayman Island:
19°20'’N, 81°10’W, 15.ii.1960 (R. A. Lewin, MCZ).
CUBA Oriente: Banes, 20°58'N, 75°43’W, 2.viii.1955
(A. F. Archer, AMNH); Ensenada Nispero, Ciudamar,
19°58’N, 75°52’W, 9.xi.1945 (P. Alayo, AMNH); Jur-
agua, 19°56’N, 75°40'W, 1.x.1955 (P. Alayo, AMNH);
Santa Fé [P?], 20°22'N, 75°53’W (A. F. Archer,
AMNH). DOMINICAN REPUBLIC Barahona: Pla-
ya Los Patos, 17°58'N, 7I°1LL'W, 31L.viii.1976 (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.1934 (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, 11.xi.1967 (A.
M. Chickering, MCZ); Palisadoes Area, 17°56'N,
76°46’W, 1.xi.1957 (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, 1987: 145, figs. 16-21, 6,
2. 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-
bo
bo
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
length 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 patella
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 mm, 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
8 © Females eX)
@ Males °
Length of spider (mm)
2
-80 = -40 0 40 80 120 160 200
Collection day (since Jan 1)
Figure 14. The length of mature M. desenderi 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 (©), stippled regression line (fF = 40%);
males (@), solid regression line (f° = 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 unlike 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 examined 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
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 January 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.ii1.1899 (AMNH), 21.iii.1899 (AMNH),
23.iii.1899 (AMNH); Archipiélago 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.1935 (Exline-Peck, CAS); Barrington Island,
0°49'S, 90°4’W, 1.viii.1929 (H. H. Cleaves, CAS);
Campion, nr. Floreana (Santa Maria), 1°15'S,
90°27'W, 1.vi.1981 (Y. Lubin, MCZ): Charles Island,
1°17'S, 90°26'W, 10.v.1899 (AMNH): Indefatigable
Island, 0°38'S, 90°23'W, 27.iv.1899 (AMNH),
18.vi.1929 (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.i1.1899 (AMNH),
20.11.1899 (AMNH), 20.iii.1899 (AMNH): Isla Bin-
dloe, O0°19'N, 90°29’W, 20.vi.1899 (AMNH): Isla
Hood, 1°23’S, 89°39’'W, 18.v.1899 (AMNH),
26.v.1899 (AMNH); Isla Pinta, S Coast, 0°35’N,
90°44'W, 25.v.1964 (D. Q. Cavagnaro, CAS); Isla Pla-
za, 0°35'S, 90°9’W, 7.iii.1970 (R. Silberglied, MCZ),
26.xi1.1973 (Y. Lubin, MCZ); Isla Santa Cruz, Acade-
my Bay, 0°44'30"S, 90°17'30"W, 13.iii.1970 (R. Sil-
berglied, MCZ); Isla Santa Cruz, E slope, 0°38’S,
90°23'W, 16.iv.1964 (D. Q. Cavagnaro, CAS); Isla
Santa Cruz, Estacion Cientifica Charles Darwin,
0°44'S, 90°18’W, 24.1.1964 (D. Q. Cavagnaro & R. O.
Schuster, CAS), 12.ii.1964 (Cavagnero & Schuster,
CAS), 3.xi.1973 (Y. Lubin, MCZ), 24.xi.1973 (Y. Lu-
bin, MCZ), 25.xi.1973 (Y. Lubin, MCZ); Isla Santa
Fé, S coast, 0°50'S, 90°4’W, 30.i1.1983 (Y. Lubin,
MCZ); James, 0°16'S, 90°42’W, 22.iv.1899 (AMNH);
METEPEIRA ° Piel 23
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.1970 (R. Silberglied,
MCZ); Tower Island, Darwin’s Bay [?], 0°17’N,
89°59’W (AMNH): Tower Islands, 0°20’N, 89°58’W,
7.iv.1925 (N.Y. Zoological Society, AMNH); W coast
of Albemarle Island, 0°11'’N, 91°21'’W, 9.iii.1935
(AMNBH).
3. Metepeira grandiosa grandiosa
Chamberlin & lvie
Figures 22-28, 321; Map 2
Metepeira grandiosa Chamberlin and Ivie, 1941: 17,
figs. 24-26, 2. Female holotype from Ben Lo-
mond, California, USA in the AMNH, examined.
Metepeira palomara Chamberlin and Ivie, 1942: 72,
figs. 200-204, 2, ¢. 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, 2, 6.
Note. Levi (1977) opted to collapse M. palustris,
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 palustris 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 median 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 o.7.) metatarsuseo 2.) tarsus lle
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
(Bigss, (ela):
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 fascicula-
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, 1.vii.1977 (C. E. Griswold, CAS).
USA Arizona: Sycamore Canyon [?] Santa Cruz Co,
31°28'N, 110°42'W, 9.ix.1978 (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 lvie
Figures 29-35, 325; Map 2
Metepeira dakota Chamberlin and Ivie, 1942: 73, figs.
205-207, 2, 5. 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 the AMNH, examined.
Metepeira grandiosa alpina:—Levi, 1977: 212-214,
figs. 99, 100, 106-111, 2, 3. Brignoli, 1983: 276.
Note. 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 to
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.
METEPEIBA ° Piel 25
grandiosa alpina
(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 lvie (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
LYON ° vigilax
= x rectangula
& ° cajabamaba
j=
se
1000
>
x
zal
3
2
So}
— 100
a
-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 (©), M. rectangula [%], 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 somewhat fatter (compare Fig. 31
with 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. grandiosa
grandiosa (Fig. 22), it is thicker and with
a sharper L-shape bend instead of a more
curved 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. je
Gertsch, AMNH). San Luis Potost: Charcas, moun-
tain side, 23°8’N, 101°7’W, 7.vii.1934 (MCZ). USA.
Colorado: Cimarron, 38°27'N, 107°33’W, 21.vii.1959
(H. W. & L. Levi, MCZ); Hayden Creek, Sangre de
Cristo Mtns., 38°25'N, 105°35’W [P], 11.vii.1961 (H.
W. & L. Levi, MCZ). Utah: SE shore of Bear Lake,
41°59'N, 111°20'W, 3.vii.1978 (G. F. Knowlton,
MCZ).
Metepeira vigilax Group
Spiders in the M. vigilax group (Mete-
peira cajabamba, Metepeira glomerabilis,
Metepeira vigilax, Metepeira rectangula)
are characterized by large, sclerotized epi-
gynal openings, not unlike a snail’s 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.ix.1955, W Weyrauch, in CAS. The specific
name is a noun in apposition after the locality.
Description. Female paratype from Ca-
jabamba, Cajamarca, Peru. Carapace dark
brown, light around eyes with lateral pos-
terior extensions (Fig. 43). Proximal two-
thirds of femora white, remainder dark
brown. 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 medians 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-
~
METEPEIRA ® Piel 2
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. cajabamba 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. cajabamba differs Som 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. cajabamba
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 cajabamba 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.
cajabamba (Fig. 43) as compared to M.
glomerabilis (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. cajabamba
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
Sho
Records Examined. PERU Ancash: Callejon de
Huaylas, 9°10’S, 77°45'W, 15.viii.1988 (V. and B.
Roth, CAS). Cajamarca: Cajabamba, 12.427, 7°37’S,
78°3’W, 25.ix.1955 (W. Weyrauch, CAS). La Libertad:
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). Lima: 20 km E An-
cén, 11°47'S, 76°59'W, 1.xi.1953 (W. Weyrauch,
CAS); 23 mi N Pativilea, 10°42’S, 77°47'W, 15.i.1955
(E. I. Schlinger and E. S. Ross, CAS); 5 km NW
Chilca, 12°29'S, 76°48'W, 12.ix.1954 (E. I. Schlinger
and E. S. Ross, CAS): between mouths of Rio Chillon
and Ancon [?], 11°54'S, 77°7'W, 5.viii.1953 (M.
Koepcke and Koepcke, MUSM); Cajacay [?], Rio
Fortaleza, 10°40’'S, 77°52’'W, 6.iii.1956 (W. Weyrauch,
CAS); Canta, Rio Chill6n, 11°28'12”S, 76°37'23’"W,
12.v.1951 (W. Weyrauch, CAS); Cerro Caracoles,
12°23'S, 76°45'W, 15.ix.1951 (W. Weyrauch, 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.1986 (D. Silva, MUSM).
6. Metepeira glomerabilis (Keyserling)
Figures 37, 46-52, 314; Map 3
Epeira glomerabilis Keyserling, 1892: 154, fig. 113,
?, 6. 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, pl. 19, fig. 10,
?. 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 glomerabilis:—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, lighter 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 Pufapi, 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. glomerabilis 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
(compare Figs. 38, 46 with Figs. 53, 60).
Metepeira glomerabilis differs from M. ca-
jabamba 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.
glomerabilis (Fig. 51), compared to M. ca-
jabamba (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 from sea level
to 4,000 m.
Records Examined. BOLIVIA Chuquisaca: Mon-
teagudo, 19°49'S, 63°59'W, 24.xii.1984 (L. E. Pefia,
AMNH). BRAZIL Espirito 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.1947 (J. C. Carvalho, MNBR)J):
Fazenda Cervo, Trés Lagoas, 20°48’S, 51°43'W,
18.ix.1964 (Exp. Depto. Zool., MZSP); Utiariti,
13°2'S, 58°1L7'W, 15.viii.1961 (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.1970 (F. M. Oliveira, AMNH); Peti Forest Res.
[P] Santa Barbara, on bushes in cerrado, 19°56'S,
43°24'W, 28.vili.1986 (R. L. C. Baptista, MZSP). Pa-
METEPEIRA ° Piel 29
@ Peru, Ecuador, Colombia
¢ Brazil, Paraguay, Bolivia
Estimated Elevation (m)
tN
=}
6
6
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: 100 = April 9; 300 = October 27; 500 =
May 15 of the following year.
Symbols: Peru, Ecuador, Colombia [Ml]; Brazil, Paraguay, Bo-
liva [@].
rand: Praia do Leste, Paranagua, 25°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. P. 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.iv.1985 (R. L. C. Baptista, MZSP). Rio
Grande do Norte: Fazenda Canaan [?], Macaiba,
5°51'S, 35°21'W, 15.ix.1951 (M. Alvorenga, MZSP).
Rondo6nia: Vila Rondonia, 10°52'S, 61°57'W, 9.ii.1961
(Pereira & Machado, MZSP). Santa Catarina: Nova
Teutonia, 27°3'S, 52°24’W, 12.v.1949 (SMF). Sdo
Paulo: Barueri, 23°31'S, 46°53'W, 13.iii.1966 (K. Len-
ko, MZSP); Campos do Jordao, 22°44'S, 45°35’W,
15.iii.1945 (Wygod, MZSP); Instituto Oceanografico,
Ubatuba, 23°26'S, 45°4'W, 15.v.1967 (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, 6.1.1985 (R. L.
C. Baptista, MZSP). COLOMBIA Cundinamarca:
Sabana de Bogota, 4°43’N, 74°10'W, 10.xii.1990 (C.
Valderrama, CV). ECUADOR Guayas: Guayaquil,
2°10'S, 79°54'W, 18.ii1.1942 (H. E. F & DL. E. FE
CAS), 22.iii.1942 (Landis, CAS). Pichincha: 15 mi N
Quito, 0°0'N, 78°30’W, 23.ii.1955 (E. I. Schlinger &
E. S. Ross, CAS). Tungurahua: Bafios, 1°24'S,
78°25'W, 15.iv.1939 (W. C. Macintyre, MCZ); Pujiapi,
1°22'S, 78°28'W, 19.vi.1943 (D. L. F. & H. E. F
CAS). PARAGUAY Alto Parand: 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.1942 (W. Wey-
rauch, AMNH). Lambayeque: 10 km S_ Chiclayo,
7°59'S, 77°17'W, 19.iii.1951 (EF. S. Ross & Michel-
bacher, CAS). Piwra: Cerro Prieto, La Brea, 4°41'S,
81°6'W (CAS); Higueron (Las Lomas) [?], 4°19’S,
80°26’W, 29.vii.1941 (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.1984 (J. Coddington,
USNM).
7. Metepeira vigilax (Keyserling)
Figures 36, 53-59, 327; Map 4
Epeira vigilax Keyserling, 1893: 211, fig. 156, ¢.
Male holotype from Taquara, Rio Grande do Sul,
Brazil, in BMNH, examined.
Araneus vigilax:—Petrunkevitch, 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
row 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
medians and anterior medians 1.2, anterior
laterals 1.2, posterior laterals 1.1. Anterior
median eyes separated by 1.8 diameters,
posterior median 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
223. tOUGt oe
Male from 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 macro-
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 medians 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 perpendicular 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
METEPEIRBA ° Piel 31
lye vigilax
ju PS (7)
Figures 38-45. 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
January, 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. 1989 (L. E. Pefia,
AMNH). Santa Cruz: Estaci6n 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,
51°9'’W, 14.vi.1964 (Celia Valle, MZSP). DOMINI-
CAN REPUBLIC Azua: El] Puerto, Majagual and
Peralta, 18°34'N, 70°47'W, 10.xi.1979 (E. Marcano,
MNSD). Prov. Trujillo Valdez: W Bani, 18°17'N,
70°22'W, 8.viii.1958 (A. F. Archer & E. de Boyrie
Moya, AMNH). HAITI Departement de L’Ouest:
Kenscoff, 18°27'N, 72°17'W, 15.xii.1929 (J. C. Myers,
AMNH), 17.iv.1935 (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 mostly 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). 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 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
Plesfountiags ule
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 with black
spots at base of setae. Femur 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).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.2, posterior laterals 1.1. Anterior median
eyes separated by 1.7 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,
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
Bode
Diagnosis. Female M. rectangula differ
from the other species in the M. vigilax
group (M. vigilax, M. glomerabilis, M. ca-
jabamba) 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 F igs.
40, 48, 55). Males differ from other spe-
cies in the group by having an embolus
that is relatively larger than that of M.
glomerabilis and M. cajabamba, 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 tiel dorsal fo-
lium has the lightest coloration (Fig. 65).
Variation. 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-
uary and February, 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.1954 Gi M. Viana,
MACN). Mendoza: Mendoza, 32°53'S, 68°49'W,
30.iii. 1965 (H. W. Levi, MCZ). CHILE Bro-Bio: 4 km
E road to Pinto, 36°42'S, 71°53’W, 4.i1.1976 (B. Mo-
reno, AMNH): Road to Pemuco, Cruce del Carmen,
36°56'S, 72°4’W, 10.1.1976 (G. Moreno, AMNH). Co-
quimbo: Illapel: Salamanca: Fundo Tahuinco,
31°44'S, 71°5’W, 30.iv.1946 (R. Doneso, AMNH).
Malleco: Angol, 37°48’S, 72°43'W (D. S. Bullock,
ie)
ee)
METEPEIRA ° Piel
CAS), 10.iii.1945 (E. A. Chapin, USNM). Maule: 10
km S Curico, 35°4'S, 71°14’W, 15.iii.1968 (L. E.
Pena, MCZ); Cordillera de Parral [?], 36°9’S,
71°50'W, 25.11.1956 (L. E. Pena, IRSNB): Linares,
35°51'S, 71°36'W (L. E. Petia, IRSNB): Miraflores,
Pedag. [?], 35°55’S, 71°39’W (Toro, AMNH).
O’Higgins: Fundo Millahue, Cunaco, 34°36'S,
71°16'W, 30.iv.1961 (AMNH). Regién Metropolitana:
Melipilla, 33°42’S, 71°13'W (L. E. Pefia, IRSNB).
Valparatso: Casablanca, 33°19'S, 71°25'W, 15.ii1.1955
(Edwin Reed, AMNH).
Metepeira labyrinthea Group
Levi (1977) described the M. labyrin-
thea group very 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. lab yinthea group and M. gosoga
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, 6, 2. Male holotype from Mexico City,
Mexico, in BMNH, examined. Roewer, 1942: 868.
Epeira labyrinthea grinnelli Coolidge, 1910: 281, °.
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, @. 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 synonymy.
Metepeira grinnelli:—Levi, 1977: 198, figs. 21-27, 6,
2. 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, other
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. lacan-
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 dimorphism 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
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, Mental: 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, ribeak 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 Aguascalientes: Hwy
45, 5.3 mi N Aguascalientes, 21°57'N, 102°17’W,
7.ix.1967 (R. E. Leech, REL). Baja California Sur:
Sierra Laguna, 17 air mi ENE Todos Santos,
93°34'N, 110°0'W, 15.xii.1979 (C. E. Griswold, CAS).
Chihuahua: 22.4 mi S Mifiaca, 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.1947 (W. le
Gertsch, AMNH):; 11 km W Suchil, 23°35'N,
104°5'W, 5.ix.1984 (W. Ile Pulawski, CAS): 6 mi NE
El Salto, 23°19’N, 105°50’W, 11.viii.1947 (W. iE
Gertsch, AMNH); Las Puentes [?], 26°49'N,
106°2'W, 23.vii.1947 (W. J. Gertsch, AMNH): Oti-
napa, Q4°11'N, 105°2'W, 12.viii.1947 (W. Je Gertsch,
AMNH): Palos Colorados, 24°2'N, 104°54’W,
5.viii.1947 (W. J. Gertsch, AMNH); Providencia,
26°44'N, 105°56’W, 24.viii.1947 (A. M. Davis,
AMNH); SW Durango, 923°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.1964
(Jean & Wilton Ivie, AMNH); 6.3 mi NW Leén,
91°13'N, 101°43’W, 6.ix.1967 (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.ix.1976
(C. E. Griswold, 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.
Born, MCZ); 4 mi N Tizayuca, 19°54'N, 98°59'W,
20.xi.1946 (E. S. Ross, CAS); 41 km N Zimapan,
20°54'N, 99°13’W, 10.viii.1991 (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.1957 (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.1948 (H. B. Leech, CAS); Charco
Ondo, 30 km W Ojuelos, 21°47’N, 101°53’W,
25.ix.1945 (H. Wagner, AMNH). México: Ixtapan de
la Sal, 18°50’N, 99°41'W, 24.viii.1946 (H. Wagner,
AMNH): Nevado de Toluca, 19°18’N, 99°44’W,
8.iv.1979 (George Uetz, MCZ); San Juan Teotihu-
acan, 19°41'N, 98°52'W, 4.xi.1939 (C. M. Bogert &
H. E. Vokes, AMNH); Tenancingo, 18°58'N,
99°36'W, 6.ix.1946 (H. Wagner, AMNH), 1.x.1946
(H. Wagner, AMNH), 15.x.1946 (H. Wagner,
AMNH); Tenango del Valle, 19°7'N, 99°33’W,
25.vili.1946 (H. Wagner, AMNH), 27.viii.1946 (H.
Wagner, AMNH); Teotihuacdn, 19°41'N, 98°52'W,
31.viiil1959 (A. F. Archer, AMNH); Tepotzotlan,
19°43'N, 99°13’W, 26.x.1982 (George Uetz, MCZ),
21.11.1983 (George Uetz, MCZ), 5.x.1983 (George
Uetz, MCZ): Toluca, 19°18'N, 99°44’W, 10.viii.1978
(George Uetz, MCZ), 1.viii.1986 (George Uetz,
MCZ); Toluca, at bottom of mountain near Parque
Cierra Morelos, 19°18’N, 99°44'W, 10.viii.1978
(George Uetz, MCZ); Toluca, E of town on Paseo
Tollocan [P], 19°18’N, 99°42’W, 23.x.1982 (George
Uetz, MCZ). México D. F.: 19°25'N, 99°10'W,
12.x.1940 (H. Wagner, AMNBH), 28.xii.1940 (R. H.
Crandall, AMNH), 15.ix.1943 (H. & D. Frizzell,
AMNH): Contreras, 19°18’N, 99°17'W, 4.xii.1944 (H.
Wagner, AMNH), 15.ix.1965 (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.1941 (H. Wagner, AMNH): EI Xitle, 18°61'N,
99°17’W [P], 12.viii.1942 (C. Tellez, AMNH): Haci-
enda Cérdoba, 19°26'N, 99°10’/W [?], 29.x.1944 (H.
Wagner, AMNH); Ouieros, 18°62’N, 99°17’W [?],
5.vii.1943 (M. Cardenas, AMNH): Mexico City,
19°25'’N, 99°10’W, 1.xi.1941 (C. Velo, AMNH),
25.ix.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. 1909 (AMNH); Petregal [?], 18°60'N,
99°17'W, 1.xii.1943 (AMNH); Rancho Cérdoba,
19°27'N, 99°10'W, 29.x.1944 (H. Wagner, AMNH):
Tlaplan, 19°17'N, 99°10'W, 7.viii.1991 (W. H. Piel &
G. S. Bodner, MCZ). Michoacan: 25 mi W. La Barca
nr. Lago de Chapala, 20°17’N, 102°34’W, 11.ix.1976
(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.1967 (R. E. Leech, REL); Hwy 15, 9.5 mi W.
Morelia, 19°42’N, 101°16’W, 18.viii.1967 (R. E.
Leech, REL); Lake Chapala, NW of Cojumatlan,
20°10'N, 102°53’'W, 7.ix.1966 (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.1941 (H.
Wagner, AMNH), 18.xi.1946 (M. G. Bradt, AMNH):
Cuernavaca/Tepotzotlan, interchange between I-95 &
115, 18°55'N, 99°13/W, 7.viii.1978 (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,
1ll.x.1994 (W. H. Piel, MCZ): S of Huitzilac,
19°0'29"N, 99°15’'50’W, 16.x.1994 (W. H. Piel, MCZ).
Puebla: 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 Potost: 3 km W. Pilares, 21°55'34’N,
100°48’6"W, 21.x.1994 (W. H. Piel, MCZ): Cuidad del
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.1959 (B. A. Branson,
AMNH): Hermosillo, 29°4'N, 110°55’W, 20.ix.1952
(B. Malkin & V. E. Thatcher, AMNH): Sierra Man-
zanal, 30°50'N, 110°10’W, 14.ix.1976 (Roth &
Schroepfer, MCZ). Tlaxcala: Huamantla, 19°19'N,
97°56'W [P], 15.vii.1981 (C. Gold, CAS). Veracruz:
15 mi W. Banderilla, 19°39'N, 97°8’W, 31.x.1973 (S.
GC. 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.1947 (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, Chiricahua
Mtns., 31°35’N, 109°14'W, 20.viii.1976 (V. Roth,
MCZ). California: 26 mi W. Santa Rosa on Hwy 116,
38°31'N, 123°4'W, 19.ix.1976 (M. E. Thompson,
MCZ): Mendocino, 39°18’N, 123°48'W, 18.viii.1959
(W. J. Gertsch, MCZ); Monterey, 36°36'N, 121°54'W,
l.ix.1949 (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
Holotype. 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 white 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-
METEPEIRBA ° Piel 37
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
Oui tountin 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 femur 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 more 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 specimens
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.viii.1976 (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.1947 (H. Wagner,
AMNH); San Cristobal de las Casas, 16°45’'N,
92°38'W, 22.vii.1947 (C. 6 ME 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 morphology.
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.,
le i, eld, IMI, 702). The embolus is
fick and Thee a large, prominent, distal
apophysis that Pees under the terminal
apophysis (Figs. 84, 92, 100, 110, 121).
11. Metepeira nigriventris (Taczanowski)
Figures, 84-91, 310; Map 5
Epeira nigriventris Taczanowski, 1878: 151, fig. 6, °.
Bovalen lectotype from Lake Junin, Peru, in PAN,
type lost. Keyserling, 1893: 217, fig. 161, 2, d.
Araneus nigriventris:—Chamberlin, ‘1916: 248. Bon-
net, 1955: 550.
Metepeira nigriventris:—Chamberlin and Ivie, 1942:
TA, figs. 211-214, 2, 6. 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). Sternum 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 De : 8 di-
ameters, posterior median eyes by 1.2, an-
terior iravetienn 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 I
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
ae
au °°
nigriventris
calamuchita
(13)
Figures 84-91. Metepeira nigriventris (Taczanowski) (sp. 11; 11°25'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, posterior.
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, epigynum,
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 likely sister species, M. tara-
paca (Fig. 94). The overall shape of the
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 slimmer and more
graceful than those of other species in the
M. nigriventris species group (compare
Fig. 84 with Bigs O27 MOOI UO! IDI).
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 specimens
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.1970 (L. E. Pefia,
MCZ). BOLIVIA La Paz: 45 mi S La Paz, 17°9'S,
67°36'W, 25.11.1951 (E. S. Ross & Michelbacher,
CAS); 70 mi S La Paz, 17°30'S, 67°36’W, 25.ii1.1951
(E. S. Ross & Michelbacher, CAS): La Paz, Avenida
Sport Club, 16°30’S, 68°9’W, 4.i1.1959 (A. M. Nadler,
AMNH); La Paz, in garden of house, 16°30'S,
68°9'W, 15.iv.1959 (R. Walsh, AMNH); Lake Titicaca,
Copacabana, Yampupata, & Isla del Sol, 16°10'S,
69°S’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.1958 (R. Walsh, AMNH);: Tiahuanaco,
Puma Puerto Ruins, 16°33'S, 68°42'W, 1.ii.1973 (Ann
Moreton, MCZ). Oruro: 6 km N Challapata, 18°51’S,
66°47'W, 23.11.1951 (E. S. Ross & Michelbacher,
CAS); Gorge Uhuschlucht, near Oruro, 17°59’S,
67°9'W, 7.ii.1954 (Forster & Schindler, ZSM). Potos?:
Villazon, 22°6'S, 65°36'W, 30.xii.1984 (L. E. Pefia,
AMNH). PERU Apurimac: Chincheros, 13°30'48’S,
73°42'47’"W, 12.xii.1980 (C. Gold, CAS); Puna near
Abancay, 13°38'2"S, 72°52'52”/W, 15.xii.1947 (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, 1.xi.1985 (D. Silva, MUSM).
Cusco: Cheqquerec, 13°23'S, 72°8'W, 2.ix.1993 (J.
Ochoa Camara, MCZ): Cusco, 13°31'6"S,
71°58’41"W, 8.viii.1965 (P. & B. Wygodzinsky,
AMNH). Junin: 8 mi W Tarma, 11°25'S, 75°48’W,
6.11955 (E. I. Schlinger & E. S. Ross, CAS); Cochas
Bajo, 11 km W Tarma, 11°25'21"S, 75°46'1L1”"W,
27.iii.1988 (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.iii1.1988 (J. Palmer,
MCZ); Huancayo, 12°4'S, 75°14'W, 15.vi.1947 (W.
Weyrauch, AMNH); Oroya, 11°32'S, 75°54'W,
12.iv.1914 (M. P. Anderson, AMNH). Lima: Bosque
de Zarate, 11°53'S, 76°27'W, 18.i.1981 (J. Francke,
MUSM). Puno: 10 mi N Ayaviri, 14°45'S, 70°35’W,
1.iii1.1951 (E. S. Ross & Michelbacher, CAS); Cama-
cane, 15°55'S, 69°50’W, 20.xi1.1955 (L. E. Pefia,
IRSNB); Isla Taquili, Lago Titicaca, 15°46’S,
69°41'W, 23.xii.1980 (C. Gold, CAS); Juli (col. Chu-
cuito), 16°13'S, 69°27'W, 7.xi.1952 (F. Blancas,
MUSM); near Chucuito, Lago Titicaca, 15°50'S,
69°48'W, 10.iii.1953 (M. Koepcke, MUSM); Puna,
Lake Titicaca, 15°50'S, 70°2'W, 15.vi.1947 (W. Wey-
rauch, AMNH); Puno, 15°50’S, 70°2’'W (Soukup,
AMNH); Yunguyo, downtown plaza, 16°15'S, 69°5’W,
31.41.1973 (Ann Moreton, MCZ).
12. Metepeira tarapaca new species
Figures 92-99, 305; Map 7
Holotype. Male from Quillagua, Antofagasta, Chile,
4.ii.1965, L. E. Pefia, 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
Estimated elevation (m
0 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 white 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 lateral eyes, lateral
eyes separated by 0.1 their diameters. To-
tal length 4.4 mm. Carapace 2 mm long,
1.6 wide. First femur 3.3 mm, patella and
tibia 3.5, metatarsus 3.3, tarsus 1. Second
patella and tibia 2.8 mm, third 1.4, fourth
Dye
Diagnosis. Unlike M. nigriventris or M.
METEPEIRBA ° Piel 4l
karkii, M. tarapaca has a white mark on
the sternum (Figs. 97, 99). The epigynum
is close to that of M. nigriventris, except
that it is less robust (compare Fig. 93 with
Fig. 85), and the membrane just over the
openings cannot be seen behind the thin-
ner scape (compare Fig. 86 with Fig. 94).
Unlike M. calamuchita, the epigynum of
M. tarapaca does not widen posteriorly
(compare Fig. 94 with Fig. 102); unlike M.
karkii, the posterior epigynal lobes are not
swollen (compare Fig. 94 with Fig. 123);
unlike M. galatheae, the black sclerotized
circles behind the epigynal openings are
larger and not shifted posteriorly (compare
Fig. 94 with Figs. 118-120). The segment
of the embolus between the basal and dis-
tal embolic apophysis is relatively thicker
than in other members of the M. nigriven-
tris species group (compare Fig. 92 with
Figs. 84, 100, 109, 121). The terminal di-
vision of the male palp in M. tarapaca
shows a distal embolic apophysis that dif-
fers in shape from that of other species
(compare Fig. 95 with Figs. 87, 103, 117,
124).
Variation. Average body length of 22 fe-
males examined 7.4 mm, range 5 to 11.5
mm. Average body length of six males ex-
amined 5.9 mm, range 4.4 to 7.3 mm. The
dorsum and venter are often darker than
the holotype. Many lack the flanking lines
and the U-shape on the venter; some at
higher elevation have a much reduced
white line on the sternum.
Natural History. Mature specimens
have been collected throughout the year,
especially between January and April (Fig.
305). V. Roth (vial label) notes that these
spiders live in a social colony. M. Roy (per-
sonal communication) reports that colo-
nies can reach 200 individuals. Median el-
evation, 2,800 m.
Dicom Moderately high altitudes
in northern Chile and southern Peru (Map
0:
Records Examined. CHILE Antofagasta: 7 km S
Toconao, 23°11'S, 68°1'W, 25.xii.1988 (V. & B. Roth,
CAS); Aguas Blancas [?] (= 24°11’S 69°51’W), To-
conao, 23°11'S, 68°L’W, 11.x.1955 (L. E. Pefia,
42, Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
IRSN); Antofagasta, 93°39'S, 70°24'W, 15.xi.1975 (L.
E. Pena, AMNH): Quillagua, 21°39'S, 69°33'W,
431.1965 (L. E. Pefia, MCZ), 211.1973 (W. C. Sedg-
wick, MCZ); Rio Loa, 25 km S Quillagua, 21°54’S,
69°33'’W, 20.viii.1966 (E. Schlinger & M. Irwin,
CAS). Elloa: Thermo Puritania, 35 km N San Pedro
de Atacama, 22°37'S, 68°13’W, 25.xii.1988 (V. & B.
Roth, CAS). Tarapaca: Arica Timar, Alociado [?],
18°45'S, 69°42’W, 22.ii1.1973 (N. Hichins, AMNH);
Ariquilda, 19°38'S, 69°32’W, 29.iv.1969 (L. E. Pena,
MCZ): Canchones, 20°27'S, 69°37'W, 29.i1.1973 (W.
C. Sedgwick, MCZ); Chapiquilta, 19°18’S, 69°25'W,
6.vi.1968 (L. E. Petia, MCZ): Chiapa, 19°32’S,
69°13’W, 24.iv.1969 (L. E. Pena, MCZ); Pica,
20°30'S, 69°21'W, 26.i1.1973 (W. C. Sedgwick, MCZ);
Poroma, Quebrada de Tarapaca, 19°52’S, 69°11'W,
20.iv.1969 (L. E. Pefia, MCZ); Quisama, 19°19’S,
69°28'W, 5.vi.1968 (L. E. Pefia, MCZ). PERU Are-
quipa: Arequipa, 16°23'56"S, 71°32’6"W, 15.vii.1955
(A. Meza, MZSP); Chiguata, near Arequipa, 16°24’S,
71°24'W, 15.ii.1948 (W. Weyrauch, MUSM).
13. Metepeira calamuchita new species
Figures 100-107, 308; Map 5
Holotype. Male from Calamuchita, Cordoba, Argen-
tina, 15.i1.1955, J. M. Viana, in MACN. The specific
name is a noun in apposition after the locality.
Description. Female paratype from Cal-
amuchita, Cordoba, Argentina. Carapace
reddish brown, light around eyes with lat-
eral posterior extensions (Fig. 106). Leg
articles annulated distally. Femur I with
row of six or seven macrosetae on anterior
side; five on anteroventral side. Anterior
portion of dorsal folium lighter than in
most species (Fig. 106). Venter black with
wide white median line and pair of large
white spots on either side of spiracle (Fig.
107). Sternum brownish black with wide,
white median line (Fig. 107). Ratio of eye
diameters: posterior medians and anterior
medians 1.1, anterior laterals 1.4, posterior
laterals 1.3. Anterior median eyes separat-
ed by 1.6 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 3.2 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 8.9
mm. Carapace 3.4 mm long, 2.8 wide.
First femur 4.5 mm, patella and tibia 4.7,
metatarsus 4.7, tarsus 1.3. Second patella
and tibia 4.1 mm, third 2.3, fourth 3.5.
Male holotype. Carapace brown with
light, triangular, median mark pointing
posteriorly (Fig. 104). Leg articles white,
darkening brown distally. Femur I with
row of four macrosetae on anterior side;
five or six on anteroventral side. Anterior
dorsal folium mostly white, posterior most-
ly black (Fig. 104). Venter as in female
(Fig. 105). Broad white median mark on
sternum (Fig. 105). Ratio of eye diame-
ters: posterior medians and anterior me-
dians 1.1, anterior laterals 1.4, posterior
laterals 1.4. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 2.2 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 4.4
mm. Carapace 2.3 mm long, 1.8 wide.
First femur 3.8 mm, patella and tibia 3.7,
metatarsus 4, tarsus 1.2. Second patella
and tibia 2.9 mm, third 1.5, fourth 2.5.
Diagnosis. Metepeira calamuchita dif-
fers from others in the M. nigriventris spe-
cies group by the large, scooped openings,
and posteriorly widening epigynum (Fig.
102). The epigyna of M. tarapaca and M.
calamuchita both have deep scooped-out
openings, but these are angled posteriorly
in the latter (Fig. 102) and laterally in the
former (Fig. 94). The male palp of M. cal-
amuchita is easily distinguished from that
of other species by the widened (Fig. 100)
and inwardly curved (Fig. 103) “dewlap”
under the embolus.
Variation. Average body length of 13 fe-
males examined 7.7 mm, range 5 to 9.8
mm. Average body length of five males ex-
amined 5.2 mm, range 4 to 6.4 mm. Two
specimens collected near 2,700 m in Itur-
be, Jujuy, Argentina, have entirely dark
sterna. In contrast, most specimens are
found at lower elevations with white me-
dian lines on their sterna.
Natural History. Mature specimens
have been collected between October and
July, although they are probably available
throughout the year (Fig. 308). Most ele-
vations range from 150 to 1,700 m, with
one population at 2,700 m.
Distribution. Northern Argentina, at
mostly lower altitudes east of the Andes
(Map. 5).
Records Examined. ARGENTINA Buenos Aires:
Las Espadufias [?], Sierra de la Ventana, 38°9’S,
61°48'W, 15.x.1973 (Maury & Cesari, MACN); Sierra
de la Ventana, 38°9'S, 61°48'W, 15.vii.1972 (Amarril-
la, MACN). Coérdoba: C. Paz, 31°24'S, 64°31'W,
15.v.1940 (C. Marti, MACN); Calamuchita, 32°4'S,
64°33’W, 15.xii.1941 (J. M. Viana, MACN), 15.i.1955
(J. M. Viana, MACN); Mina Clareo, 31°43’S, 65°0'W,
15.iv.1973 (Stiebel, MACN). Jujuy: Iturbe, 22°59’S,
65°21'W, 22.ii.1983 (L. E. Pefia, AMNH). La Rioja:
Ilias [?], 29°5’S, 66°19’W (P. M. Gomez, MACN).
San Luis: Chosmes and Desaguadero (Mendoza),
33°24'30"S, 67°0'0"W, 14.iv.1967 (L. E. Pefia, MCZ).
Santiago del Estero: Santiago del Estero, 27°47’S,
64°16'W (AMNH), 3.iv.1965 (H. W. Levi, MCZ): 70
km W Santiago, 27°47'S, 65°25'W, 3.iv.1965 (H. W.
Levi, MCZ); Quebrachos: Sumampa, Parayacu [?],
29°22'S, 63°28'W, 15.xi.1944 (Maldonado Bruzzone,
MLBP).
14. Metepeira galatheae (Thorell)
Figures 108-120, 304; Map 7
Epeira galatheae Thorell, 1891: 53. Female holotype
from “Cobija, Bolivia,” now, Cobija, Antofagasta,
Chile, in the UZMK, examined.
Araneus galatheae:—Bonnet, 1955: 506.
Metepeira galatheae:—Levi, 1991: 179. Platnick,
1993.
Metepeira cereicola nomen nudum, female in AMNH
but no description has been found. Female man-
uscript type from Salamanca, Coquimbo, Chile,
collected by Archer on 30.iv.1961.
Note. Thorell (1891) listed the holotype’s locality
as “Cobija, Bolivia.” While Cobija, Bolivia exists
(11°02'S 68°44'W), it is an unlikely locality for the
corvette Galathea to visit on its voyage to southern
Asia because it would have meant climbing over
the Andes. Instead, the port town of Cobija, Chile
(22°33'S 70°16'W) is much more likely, especially
since this region of Chile was under Bolivian ad-
ministration throughout the period of Galathea’s
voyage, 1845-1847 (Paynter et al., 1975).
Description. Female from Chile Chico,
Aisén province, Chile. Reddish brown car-
apace with white setae, light around eyes,
lateral posterior extensions, central arrow-
shaped mark (Fig. 113). Legs light yellow,
articles annulated distally. Femur I with
row of four or five macrosetae on anterior
side; two to four on anteroventral side.
Dorsum with black and white setae. Foli-
um speckled brown with a white fleur-de-
lis that reduces posteriorly (Fig. 113). Ven-
ter brownish gray with wide median white
METEPEIRA ° Piel 43
line and pair of large white spots on either
side of spiracle. A pair of very faint white
stitching parallel to and on either side of
median line (Fig. 114). Sternum reddish
brown, sometimes with small posterior and
anterior white marks (Fig. 114). Ratio of
eye diameters: posterior medians and an-
terior medians 1.0, anterior laterals 1.2,
posterior laterals 1.2. Anterior median
eyes separated by 1.7 diameters, posterior
median eyes by 1.2, anterior median eyes
separated from anterior laterals by 3.6 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.1 their diameters. To-
tal length 7.8 mm. Carapace 3.5 mm long,
2.7 wide. First femur 3.7 mm, patella and
tibia 3.9, metatarsus 3.5, tarsus 1.2. Sec-
ond patella and tibia 3.4 mm, third 2.1,
fourth 3.
Male from Chile Chico, Aisén province,
Chile. Leg coloration as in female, except
femur I mostly reddish brown with row of
four macrosetae on anterior side; five on
anteroventral side. Carapace, dorsum, ven-
ter and sternum as in female (Figs. 111,
112). 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 1.6 diameters,
posterior median eyes by 1.2, anterior me-
dian eyes separated from anterior laterals
by 3.3 diameters of anterior lateral eyes,
lateral eyes separated by 0.4 their diame-
ters. Total length 5.3 mm. Carapace 2.7
mm long, 2.1 wide. First femur 3.9 mm,
patella and tibia 3.8, metatarsus 3.6, tarsus
1.1. Second patella and tibia 3.2 mm, third
1.8, fourth 2.6.
Diagnosis. Although the sternum of M.
galatheae varies from solid brown (Fig.
114) to brown with median white marks,
the venter of the abdomen is surprisingly
consistent. The venter has a wide brownish
gray area and a short median white line
flanked by very faint indications of parallel
lines (Fig. 114). In contrast, the flanking
lines in M. karkii thicken anteriorly (Fig.
128) and the venter of M. nigriventris is
almost completely black (Fig. 91). Despite
enormous variation in the hood of the
44 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Collection Day (since Jan 1)
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
Bto-Bio: Chillan, 8.xi.1976 (G. Moreno,
AMNH); Las Lajuelas, 11.i.1976 (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 February; 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
1s
Records Examined. ARGENTINA Buenos Aires:
Felipe Sola, 38°1'S, 62°50’W, 15.i.1944 (Prosen,
MLP); Patagones, 40°48'S, 62°59’W, 15.ii.1937 (J. M.
Viana, MACN): Sierra de la Ventana, 38°9’S,
61°48'W, 15.iii.1939 (J. C. Gario, MACN). Catamar-
ca: Mutquin, 28°19'S, 66°10’W, 15.i1.1963 (O. de Fer-
rariis, AMNH). Chubut: 15 km S Epuyen, 42°22'S,
71°21'W, 15.i.1986 (P. A. Goloboff, N. I. Platnick, &
R. T. Schuh, AMNH): 19.5 km E Shaman, 44°27'S,
70°30'W, 19.xi.1966 (E. I. Schlinger & M. E. Irwin,
CAS); 3 km N Puerto Lobos, 41°59'S, 65°6'W,
14.xii.1966 (E. I. Schlinger & M. E. Irwin, CAS); 35
km E Esquel, 42°54'S, 70°53’W, 18.xi.1966 (E. I.
Schlinger & M. E. Irwin, CAS); El Hoyo [?], 42°4’S,
71°30'W (A. Kovacs, AMNH), 10.i.1962 (Andor Ko-
vacs, AMNH); Epuyen, 42°15'S, 71°23'W, 18.xi.1962
(Andor Kovacs, AMNH); Leleque, 42°28’S, 71°6’W,
12.11.1965 (Andor Kovacs, AMNH); Los Manantiales,
METEPEIRA ° Piel 45
A 4
ee 2
BS
galatheae
(14)
karkil
(is)
Figures 110-120. Metepeira galatheae (Thorell) (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. 111, 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 karkii (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.1985 (L. E. Pefia, AMNH): N of Camarones,
Cantera, Namuncura, 44°46'S, 65°42/W, 17.xi.1985
(L. E. Pefia, AMNH): Rio Turbio, 42°13'S, 71°41’W
(Andor Kovacs, AMNH), 12.i.1962 (Andor Kovacs,
AMNH). Cordoba: 12 mi W Sampacho, 33°23’S,
64°43'W, 7.ii.1951 (E. S. Ross & Michelbacher, CAS):
Arguello, 31°21'S, 64°15'W, 15.xii.1943 (J. A. De Car-
\
lo, MACN); Calamuchita, 32°4'S, 64°33’'W,
15.xii.1940 (J. M. Viana, MACN); Sampacho, 33°23’S,
64°43'W, 7.i1.1951 (E. S. Ross & Michelbacher, CAS).
Mendoza: Between Beazley and San Rafael, 34°10'S,
67°29'W, 4.iii.1983 (L. E. Petia, AMNH); Mendoza,
32°53'S, 68°49'W, 30.iii.1965 (H. W. Levi, MCZ); Us-
pallata, 32°35'S, 69°20'W, 7.iii1.1983 (L. E. Pefia,
AMNH). Neuquén: Catan Lil, Charahuilla, 39°45’S,
46 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
70°37'W, 15.ii.1971 (O. de Ferrariis, AMNH):; Cuba
del Leon [?], 39°9’S, 70°53’W, 15.1.1975 (Maury,
MACN); Lago Alumine, 38°55’S, 71°9’W, 15.i.1976
(O. de Ferrariis, AMNH); Zapala, 38°54’S, 70°4’W,
15.i.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.ii.1959 (J.
R. Navas, MACN); El Bolson, 41°58’S, 71°31'W,
1.ii.1961 (A. Kovacs, AMNH), 17.x.1961 (Andor Ko-
vacs, AMNH); El Bolson, 41°58’S, 71°35’'W, 2.ii1.1965
(Andor Kovacs, AMNH): General Roca, 39°3'S,
67°32'W, 15.ix.1964 (Bachmann, MEG). Salta: Maury
[P], 24°40'S, 65°45'W, 15.1.1975 (MACN). San Juan:
10 km N Matagusanos, 31°10’S, 68°38’W, 13.i.1983
(L. E. Petia, AMNH). San Luis: INTA Experimental
Station, E of Villa Mercedes, 33°40'S, 65°27'W,
8.xii. 1967 (C. R. Ward, CAS). Santa Cruz: 2.4 km S$
Fitz Roy, 47°2'S, 67°15'W, 12.xii.1966 (E. I. Schlinger
& M. E. Irwin, CAS). Tucumdn: San Miguel de Tu-
cuman, IML gardens, 26°49'S, 65°13'W, 19.xii.1979
(L. A. Stange, FSCA). BRAZIL Mato Grosso: Campo
Grande, 20°27'S, 54°37'W, 7.ii.1952 (M. Alvarenga,
MZSP). CHILE Aconcagua: W end tunnel, 85 km S
Hlapel, 32°49'S, 71°7'W, 29.xi.1950 (E. S. Ross &
Michelbacher, CAS). Aisén: 8 km W Chile Chico,
46°33'S, 71°57'W, 22.xi.1966 (E. I. Schlinger & M.
E. Irwin, CAS); Chile Chico, near lake, 46°33’S,
71°43'W, 21.xi.1966 (E. I. Schlinger & M. E. Irwin,
CAS). Antofagasta: 6 km N Muelle de Piedra, N Tal-
tal, 25°21'S, 70°30’W, 4.ii.1942 (Junius Bird, AMNH);
Caleta Hueso Parado, Taltal, 25°22’S, 70°2S’W,
1.ii.1941 (Junius Bird, AMNH); Cobija, 22°33’S,
70°16'W (ZMUC); Quebrada Paposo, 25°2'S,
70°27'W, 3.ii.1989 (L. Stange, FSCA). Araucania: Pe-
mehue [?], 38°3'S, 71°43’W (L. E. Pefia, IRSNB):
Villarrica, 36°16'S, 72°13'W, 25.xi.1963 (G. F. Ed-
munds, AMNH). Atacama: 50-60 km S Copiapo,
2751'S, 70°20'W, 24.viii.1966 (E. I. Schlinger & 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.vi.1968 (L. E. Pefia, MCZ). Bio-Bio: 4 km E road
to Pinto, 36°42’S, 71°53’W, 4.1.1976 (B. Moreno,
AMNH); Chillén, 36°36’S, 72°7’W, 2.i.1976 (G. Mo-
reno, AMNH), 21.ii.1978 (G. Moren, MCZ); Chillan,
in cemetery, 36°36’S, 72°7'W, 8.xi.1976 (G. Moreno,
AMNH); Cuesta de Quilmo, Chillin, 36°38'S,
72°12'W, 13.xi.1976 (G. Moreno, AMNH); El Aban-
ico, 37°20'S, 71°31’W, 30.xii.1950 (E. S. Ross &
Michelbacher, CAS); Las Lajuelas, 36°39'S, 72°8’W,
111.1976 (G. Moreno, AMNH). Concepcion,
36°50'S, 73°3'W (L. E. Pefia, IRSNB); Concepcion:
Salta de Rio Laja, 37°13’S, 72°23’W, 30.i.1951 (E. S.
Ross & Michelbacher, CAS); Nuble: 50 km E San
Carlos, 36°25'S, 71°6’W, 26.xii.1950 (E. S. Ross &
Michelbacher, CAS); Nuble: Cordillera de Chillén
[?], 36°51'S, 71°24’W, 1.ii.1947 (L. E. Pefia, IRSNB):
Rio Andalien, 36°44'S, 73°1’W, 25.iii.1979 (S. Gu-
tierrez, MCZ). Coquimbo: 20 mi E La Serena,
29°54'S, 70°56’W, 3.vii.1950 (E. S. Ross & Michel-
bacher, CAS); 5 mi N Ovalle, 30°31’S, 71°12’'W,
1.xii.1950 (E. S. Ross & Michelbacher, CAS); Banos
del Toro, 29°50'S, 70°2’W, 15.ii.1947 (L. E. Pefia,
IRSNB); Cerro Talinay, 30°50'29"S, 71°37'14”"W,
29.xi.1961 (A. F. Archer, AMNH); Cuesta las Cardas,
Ovalle Rd., 30°17'S, 71°16'W, 13.xi.1961 (R. Wagen-
knecht, AMNH); Hacienda Illapel, 31°36’S, 71°7’W,
3.x1.1954 (L. E. Pefia, IRSNB), 19.x.1966 (E. I.
Schlinger, M. E. Irwin, & L. E. Pefia, CAS); Illapel:
Salamanca: Fundo Quelén, 31°52’S,— 70°52’W,
30.iv.1961 (A. F. Archer, AMNH): La Serena,
29°54'28"S, 71°15'15"W, 15.ii.1947 (L. E. Pefia,
IRSNB); Loma de Pefiuelas, 6 km S La Serena,
29°57'S, 71°18'W, 28.xi.1961 (A. F. Archer, AMNH):
Quilacén, 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.ii.1955 (Edwin Reed,
AMNH): Rio Bueno, 40°19’S, 72°58’W (L. E. Pejia,
IRSNB): Valdivia: Neltume, 39°48'S, 71°57’'W,
23.x1.1988 (V. & B. Roth, CAS). Malleco: Angol,
37°48'S, 72°43'W, 29.i1.1951 (E. S. Ross & Michel-
bacher, CAS). Maule: Linares, 35°51'S, 71°36’W (L.
E. Pena, IRSNB), 15.1.1947 (L. E. Pena, IRSNB):
Miraflores, Pedag. [?], 35°55’S, 71°39'W (Toro,
AMNH). O’Higgins: Chépica, 34°44'S, 71°17'W,
15.xii.1947 (L. E. Pefia, IRSNB). Regidn Metropoli-
tana: 34 km W Santiago, 33°30'S, 71°25'W,
19.xii.1950 (E. S. Ross & Michelbacher, CAS); Ba-
tuco, nr. Santiago, 33°13'S, 70°47’W (Guil. Mann,
AMNH); Lampa, 33°17'S, 70°54’W, 1.v.1979 (L. E.
Pefia, AMNH). Santiago: El Golf [?], 33°30'S,
71°25'W, 9.iv.1961 (A. F. Archer & J. Aros, AMNH);
Santiago, 33°30'S, 71°25'W (L. E. Pefia, IRSNB),
1.ii.1973 (W. C. Sedgwick, MCZ). Talca: 22 mi N Tal-
ca, 35°7'S, 71°40'W, 22.xii.1950 (CAS). Valparaiso:
Concon, in cow farm, 32°55'S, 71°31'/W, 4. iii. 1962 (H.
Morales, AMNH); La Cruz, 32°53’'S, 71°16'W,
18.i.1973 (W. C. Sedgwick, MCZ); Llay-Llay, 32°51’S,
70°58’W, 20.i1.1973 (W. C. Sedgwick, MCZ): Los Mai-
tenes [P], 32°59’S, 71°15’W, 14.x.1954 (L. E. Pefial,
IRSNB); Quintay, 33°11'S, 71°42'W, 19.ii.1967 (E. I.
Schlinger, CAS); Valparaiso, 33°2'S, 71°38’W (Edwin
Reed, AMNH).
15. Metepeira karkii (Tullgren)
Figures 121-128, 303; Map 5
Araneus karkii Tullgren, 1901: 219, 259. Female ho-
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
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 medians
and anterior medians 1.2, anterior laterals
1.4, posterior laterals 1.5. Anterior median
eyes separated by 1.8 diameters, posterior
median 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 mm. 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
mark (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
sternum 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 median
eyes separated by 1.7 diameters, posterior
median 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
METEPEIRBA °® Piel AT
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. calamuchita 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 vary 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.1988 (V. & B. Roth, CAS). Neuquén: Laguna
Blanca, 39°3’S, 70°23'W, 15.iii.1959 (J. Nara,
MACN); Zapala, 38°54'S, 70°4'W, 15.i1.1958 (fe R. Na-
vas, MACN); Zapala, Laguna Blanca, 38°54'S,
70°4'W, 15.1.1959 (J. R. Navas, MACN). Rio Negro:
Cerro Alto [?], 41°8’S, 70°40’W (MACN): Coronel
Juan José Gomez, 39°2'S, 67°39'W, 15.xi.1945 (Ibarra
Grasso, MLP); Ne-Luan, 41°25'S, 68°45’W (MACN).
Santa Cruz: Laguna Calafate, Precordilfera [?],
50°55’'S, 70°9'W, 22.1.1967 (P. San Martin, MCZ): Rio
Gallegos, 51°38’S, 69°13’W, 20.i.1967 (P. San Martin,
MCZ). BOLIVIA Santa Cruz: 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.xii.1966 (E. I.
Schlinger & M. E. Irwin, CAS); Kark, 51°17'30"S,
72°32'30"W, 13.iii.1899 (FE. Nordenskidld, NRMS).
Metepeira compsa Group:
The three species in the M. compsa
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).
16. Metepeira compsa (Chamberlin)
Figures 129-140, 311; Map 8
Aranea compsa Chamberlin, 1916: 252, fig. 6, 2. Fe-
male holotype from Ollantaitambo, Cusco, Peru, in
the MCZ, examined. Bonnet, 1955: 462.
Araneus labyrintheus:—Petrunkevitch, 1926: 27. Er-
roneous synonymy.
Metepeira virginensis Chamberlin and Ivie, 1942: 70,
figs. 188-190, 2, d. Female holotype from St.
Thomas, U.S. Virgin Islands, in AMNH, examined.
NEW SYNONYMY.
Metepeira latigyna Chamberlin and Ivie, 1942: 70,
figs. 191, 192, 2. Female holotype from Porto Ale-
gre [Bahia], Brazil, in AMNH, examined. NEW
SYNONYMY.
Metepeira compsa:—Chamberlin and Ivie, 1942: 71,
figs. 193-195, @, 3.
Metepeira labyrinthea:—Bryant, 1942: 346.
Metepeira perezxi Archer, 1965: 132, figs. 14, 16, 3,
2. Male holotype from subexperiment station, Is-
abela, Puerto Rico, in AMNH, examined. Brignoli,
1983: 276. NEW SYNONYMY.
Metepeira vaurieorum Archer, 1965: 133, figs. 15, 19,
3, 2. Male holotype from Usine de Robert, Mar-
tinique, in AMNH, lost. Brignoli, 1983: 276. Lo-
pez, 1993: 10, 11, figs. 1-4, 11, 6. NEW SYN-
ONYMY.
Note. Although the type for M. vaurieorum is
lost, all examined records from Martinique (includ-
ing ones from “Usine de Robert”) belong to M.
compsa.
Description. Female from Savonet, Cu-
ragao, 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
mm, third 0.9, fourth 1.5.
Male from Savonet, Curagao, 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 mm 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 rim (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
(i)
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 specimens
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, 15.11.1941 (F. Moneos,
MACN). Chaco: Basail, 27°52'S, 59°18’'W, 18.iv.1942
(MACN). Corrientes: Paso de la Patria, 27°19’S,
58°35'W, 151.1966 (M. E. Galiano, MEG). Entre
Rios: Parana, 31°44’'S, 60°32'W (Rosenzwaig, MLP);
Salto Grande, 31°13’S, 57°56'W, 15.iii.1964 (M. E.
Galiano, MEG). Misiones: Posadas, 27°23'S, 55°53’ W,
15.ix.1963 (M. E. Galiano, MEG). Neuwquén: Piedra
del Aguila, 40°3’S, 70°5’'W, 18.vii.1978 (Mision Cien-
tifica Danesa, ZMUC). Santa Fé: Tostado, 29°14'S,
61°46'W (A. Giai, MACN). BOLIVIA La Paz: Apolo,
14°43'S, 68°31'W, 10.viii. 1989 (L. E. Pefia, AMNH).
BRAZIL Bahia: Arquipélago dos Abrolhos, 17°40'S,
38°50'W, 28.xii.1887 (U.S.F.C., 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 Azul,
16°1'S, 41°16’W, 15.xii.1970 (F. M. Oliveira, AMNH).
Pernambuco: Pernambuco [?], 8°3'S, 34°54’W,
12.ii1.1927 (SMF), 8.iii.1955 (SMF). Rio de Janeiro:
Lagomar [?], Macae, near sea, 22°23'S, 41°47'W,
17.vii.1986 (R. L. C. Baptista, MZSP). Rio Grande do
Sul: Montenegro, 29°42'S, 51°28'W, 3.xi.1977 (E. H.
Buckup, MCN); Rambo [?], 30°4'0"S, 51°11’W
(MNRJ); Sao Leopoldo, 29°46'S, 51°9'W, 14.x.1965
(Celia Valle, MZSP). Sdo Paulo: Rubiao, Jr. [?], Bo-
tucatu, 22°52’S, 48°26’W, 25.iv.1988 (R. L. C. Bap-
tista, MZSP); Sao Paulo, Guaruja, 24°0’S, 46°16’W,
24 vii.1983 (R. Sievers, AMNH). BRITISH WEST
INDIES Anegada: 18°45'N, 64°20'W, 12.xi.1966
(Harry Beatty, AMNH); center, nr. salt pond,
18°45'N, 64°20'W, 4.vi.1966 (Island Project Staff,
Univ. of Puerto Rico, AMNH): West end, 18°45’N,
64°22'W, 4.vi.1966 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Anegada Settlement: 18°45’N,
64°20'W, 5.vi.1966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Antigua: Coolidge Airport,
17°6'N, 61°51’W, 15.xi.1967 (N. L. H. Krauss,
AMNH): Devil’s Bridge, 17°5'N, 61°42'W, 30.vi.1963
(Rick & E. N. Kjellesvig-Waering, AMNH); Jolly
Beach, 17°4’N, 61°53’W, 20.ix.1963 (E. N. Kjellesvig-
Waering, AMNH); Lignum Vitae Bay: Jolly Beach [?],
17°4'N, 61°53'W, 19.vi.1968 (E. N. Kjellesvig-Waer-
ing, AMNH); Redonda Island, from webs spun be-
tween boulders on beach, 16°55'’N, 62°19'W,
10.iv.1956 (J. F. G. Clarke, USNM); Reeds Point, nr.
Jolly Beach, 17°4'N, 61°53’W, 2.vii.1963 (E. N. Kjel-
lesvig-Waering, AMNH); Saint John’s, 17°6'N,
6L°51'W, 15.viii. 1967 (N. L. H. Krauss, AMNH),
15.xii. 1967 (N. L. H. Krauss, AMNH). Dead Man’s
Chest [?]: 18°22'N, 64°34'W, 26.v.1966 (Island Pro-
ject Staff, Univ. of Puerto Rico, AMNH). East Seal
Dog: 18°30'N, 64°25'W, 7.vi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). George Dog Is-
land: 18°30'N, 64°27'W, 7.vi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH); 4 mi W Virgin
Gorda Isl, high on dry cliffs, 18°30'N, 64°27'W,
25.ii1.1979 (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.1966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Green Cay, near Tortola:
18°27'N, 64°42'30"W, 14.viii.1965 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Island O. near
Anegada Settlement: 18°45'N, 64°20’W, 5.vi.1966 (Is-
land Project Staff, Univ. of Puerto Rico, AMNH). Is-
land R. near Anegada Settlement: 18°45’'N, 64°20'W,
4.vi.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Jost Van Dyke: 18°28'N, 64°45'W,
30.viii.1965 (H. Heatwole, R. Levins & F. Mac-
Kenzie, AMNH). Large mangrove patch nr. Settle-
ment Anegado: 18°45'N, 64°20'W [?], 5.vi.1966 (Is-
land Project Staff, Univ. of Puerto Rico, AMNH). Lit-
tle Camanoe: 18°28'N, 64°33’W, 25.v.1966 (Island
Project Staff, Univ. of Puerto Rico, AMNH). Little
Jost 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.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Montserrat:
Gage’s Soufriére [?], 16°43’N, 62°10’W, 28.vii.1972
(N. L. H. Krauss, AMNH); Plymouth, 16°42’N,
62°13'W, 15.xi.1967 (N. L. H. Krauss, AMNH). Neck-
er Island: 18°33'N, 64°21'W, 6.vi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). nr. Anegada Set-
tlement: Byers Bache [P?], 18°45'N, 64°20’'W,
5.vi.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Peter Island: 18°22'N, 64°35'W, 6.vii.1965
(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.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Saint Christo-
pher Island: Basseterre, 17°18'N, 62°43'W, 6.ii.1968
(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.1965 (H. & A. Heatwole, AMNH). Tobago Is-
land: 18°27'N, 64°48'W, 2.iv.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Tortola:
18°27'N, 64°36'W, 8.vii.1958 (A. F. Archer, C. Hel-
sley, & M. Sanderson, AMNH); Beef Island, 18°27'N,
64°31'W, 21.vii.1965 (H. Heatwole, R. Levins & F.
MacKenzie, AMNH); Greater Camanoe Isl, 18°29'N,
64°32’W, 1.vii.1965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Little Thatch Island, 18°23'N,
64°42'W, 16.viii.1965 (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.1965 (H. Heatwole & R. Levins,
AMNH); Prospect Reef, S Roadtown, 18°25'N,
64°36'W, 23.iii. 1979 (K. Johnson, AMNH), 31.iii.1979
(K. Johnson, AMNH); Road to town, 18°27’N,
64°36'W, 15.vii.1972 (N. L. H. Krauss, AMNH):
Sandy Spit [?], 18°27’N, 64°36’W, 14.viii.1965 (Island
Project Staff, Univ. of Puerto Rico, AMNH). Tortola
Island: 18°27'N, 64°36'W, 15.vii.1965 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Virgin Gorda:
18°30'N, 64°24’W, 15.xi. 1966 (Harry Beatty, AMNH):
Baths & Devil’s 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);
Trellis 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.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). West Seal Dog Island: 18°29'N, 64°28'W,
7.vi.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). CHILE Tarapaca: Arica, 18°29’S, 70°20'W,
28.i.1973 (W. C. Sedgwick, MCZ); Azapa Arica,
18°31'S, 7O°1L1’W, 9.xi.1955 (L. E. Petia, IRSNB):;
Lluta, 18°24'S, 70°19’W, 12.xi.1955 (L. E. Pefia,
IRSNB); Rio Lluta, 18°24'S, 70°19’W, 12.xi.1955 (L.
E. Pena, IRSNB); Sobraya, 18°32'S, 70°9'W,
10.xi.1955 (L. E. Petia, IRSNB); Taltape, Camarones
Valley, 18°59'S, 69°47'W, 29.i1.1973 (MCZ). COLOM-
BIA Magdalena: Cabaiia “Villa Culebra,” 10 km E
Station Marta, 11°12’N, 74°7’W, 15.x.1985 (H.-G.
Miiller, SMF); Casajera [P], 11°O0’N, 74°15’W,
1.ii.1974 (J. A. Kochalka, JAK); Cienaga, 11°1’N,
74°15'’W, 30.i1.1974 GE A. Kochalka, JAK); Gaira,
11°11'N, 74°13'W, 15.xii.1975 (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.1960 (C.
& P. Vaurie, AMNH); Marie-Galante, in citrus,
15°56'N, 61°16’W, 15.iii.1977 (W. H. Whitcomb,
FSCA): Pointe-a-Pitre, [let A Boissard, 16°14'N,
61°34'W, 26.vi.1960 (C. & P. Vaurie, AMNH): Terre-
de-Haut, Les Saintes, 15°58’N, 61°35/W, 2.vii.1960
(C. & P. Vaurie, AMNH). Martinique: Ansemitian [?],
METEPEIRA ® Piel Sail
Trois-Ilets, 14°33'N, 61°2'W, 10.vi.1960 (C. & P. Vau-
rie, AMNH):; Diamant, 14°29'N, 61°2’W, 17.vi.1960
(C. & P. Vaurie, AMNH), 18.vi.1966 (C. & P. Vaurie,
AMNH); Fort de France, 14°36’N, 61°5’W,
15.xii.1950 (N. L. H. Krauss, MCZ); Pointe Ferret,
La Caravelle, 14°45'N, 60°54’W, 19.vi.1960 (C. & P.
Vaurie, AMNH); Sainte-Anne, 14°26’N, 60°53’W,
20.vi. 1966 (C. & P. Vaurie, AMNH): Usine de Robert
[P], 14°41'’N, 60°57'W, 16.vi.1960 (C. & PB. Vaurie,
AMNH). GRENADA nr. Saint Georges: 12°3'N,
61°45'W, 3.vi.1950 (Leo Isaacs, AMNH). NETH-
ERLAND ANTILLES Bonaire: Red Pond [?],
12°12'N, 68°15'W, 3.i1.1968 (B. Malkin, AMNH). Cu-
racgao: 12°11'N, 68°58'W, 13.i1.1968 (B. Malkin,
AMNH); Fuik (Oostpunt), muddauber nests, 12°4'N,
68°49'W, 20.xii.1962 (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. 1962 (H. W. & L. Levi,
MCZ): Hato, 12°11'N, 68°58’W, 28.xii.1967 (B. Mal-
kin, AMNH), 7.i.1968 (B. Malkin, AMNH): Piscadera
Baai, 12°8'N, 68°59’W, 18.xii.1962 (H. W. Levi,
MCZ), 20.xii.1962 (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.1962 (H. W. Levi & B. de Jong, MCZ): Siberié,
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. 1962 (H. W. Levi, MCZ). Sint
Eustatius: Oranjestad, 17°29'N, 62°59'W, 18.i.1968
(B. Malkin, AMNH). Sint Maarten: nr. Juliana Air-
port [P], 18°4’N, 63°4'W, 24.ii.1965 (H. Heatwole &
F. MacKenzie, AMNH). PARAGUAY Alto Paranda:
Taguarazaya [?], 25°30'S, 54°50’W (AMNH). PERU
Ancash: Huaraz, 9°32'S, 77°32'W, 6.xii. 1980 (C. Gold,
CAS). Apurimac: 35 mi E Abancay, 13°38'S, 72°22'W,
5.i1.1951 (E. S. Ross & Michelbacher, CAS); 40 mi
E Abancay, 13°38’S, 72°20'W, 4.iii.1951 (E. S. Ross
& Michelbacher, CAS); Abancay, 13°38'2"S,
72°52'53"W, 6.iii.1951 (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.1951 (E. S. Ross &
Michelbacher, CAS); Hacienda Urco, near Calea,
13°22’S, 71°54'W, 19.ix.1939 (Karl P. Schmidt,
AMNH); Huacerpay [?], 13°37'S, 72°13'W, 10.ix.1993
(J. Ochoa Camara, MCZ); Ollantaitambo, 13°15'17'S,
72°15'48"W, 15.vii.1911 (Yale Peruvian Expedition,
AMNH), 15.xii.1980 (C. Gold, CAS): Pisac,
13°25'21"S, 71°50'48’W, 13.xii.1980 (C. Gold, CAS);
Urubamba, 13°18’28"S, 72°6'55’"W, 15.i.1965 (Carras-
co, MCZ), 18.11.1965 (H. W. Levi, MCZ), 6.viii. 1987
(D. Silva, MUSM). Huanuco: Huanuco, 9°55’S,
76°14'W, 6.x.1946 (J. C. Pallister, AMNH). Ica: km
367 between Ica and Nazca, sandy semidesert,
14°24'S, 75°23'W, 22.41.1952 (W. Weyrauch, CAS):
Nazca, 14°50'S, 74°57'W, 15.iv.1951 (P. Aguilar,
CAS). Piura: Cerro Prieto, La Brea, 4°41'S, 81°6’W
(CAS); Mallares, 4°51'S, 80°46’'W, 8.vi.1941 (H. E. F.
& D. E. F, CAS), 13.vii.1941 (WH. E.F & DL ELF
52 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
CAS); N of Negritos, Parifias 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.iii.1939 (H. E. F, CAS): nr. Se-
chura, 5°33'S, 80°51'W, 4.xi.1941 (D. L. F, CAS); Ne-
gritos, 4°49'S, 81°18’W, 15.xi.1934 (H. E. F, CAS),
15.iii.1941 (H. E. F, CAS): Parifias, 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); Parifias Valley, 4°30'S, 81°8'’W,
6.iii.1939 (D. L. & H. E. Frizzell, CAS), 8.iv.1939 (D.
L. & H. E. Frizzell, CAS); Portachuelo, 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.1939 (D.
L. & H. E. Frizzell, CAS), 11.vi.1939 (D. L. & H. E.
Frizzell, CAS); Rio Chira Valley, N of Amotape,
4°53'S, 81°1L'W, 13.xi.1938 (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.1969 (P. Aguilar, MCZ); Sullaiia,
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 Punta Parifias, 4°18’S,
80°51’W, 1.1.1939 (D. L. & H. E. Frizzell, CAS).
SAINT LUCIA Castries: 14°1'N, 61°0'W, 28.vii.1963
(E. N. Kjellesvig-Waering, AMNH). TRINIDAD &
TOBAGO Tobago Island: Buccoo Bay, 11°10'N,
60°48'W, 15.viii.1965 (E. N. Kjellesvig-Waering,
AMNH); Guayaguayare. Point, 10°8'’N, 61°2'W,
14.ix.1963 (E. N. Kjellesvig-Waering, AMNH); Pi-
geon Point, 11°10'N, 60°50’W, 18.viii.1937 (E. D.,
MCZ); Salybia Bay, by shore, 10°42'N, 61°2’W,
15.ii.1972 (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.1944 (R. Montgomery, AMNH). URUGUAY Co-
lonia: Punta Gorda [?], 34°28'S, 57°51'W, 25.ii.1968
(R. Capocasale & L. Bruno, CAS). USA Puerto Rico:
Algodones Key, 18°12’N, 65°41'W, 15.x.1964 (H.
Heatwole, R. Levins & F. MacKenzie, AMNH); Bal-
neario [?] Guajataca, 18°21'N, 66°55'W, 4.vii.1958 (A.
F. Archer, AMNH): Bafios de Coamo, 17°59’'N,
67°3'W, 2.iv.1990 (H. W. & L. Levi, MCZ), 3.iv.1990
(H. W. & L. Levi, MCZ); Barranquitas, 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.iii1.1989 (H. W. & L. Levi, MCZ): Cabeza de Perro
Island, 18°15’N, 65°35’W, 16.i1.1965 (H. Heatwole &
F. MacKenzie, AMNH); Cayo Don Luis, 17°57’'N,
66°58'W, 12.i.1966 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Cayo Norte, off Culebra,
18°20'N, 65°15’W, 14.iv.1965 (H. Heatwole & F.
MacKenzie, AMNH); Cayo Pifierito, 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.1965 (Island Project Staff,
Univ. of Puerto Rico, AMNH); Cuevas de los Alfaros,
Barrio Mora, 18°29'N, 67°1'W, 20.vii.1958 (A. F. Ar-
cher & Rolle, AMNH):; Culebra Island, 18°19’N,
65°17'W, 19.vii.1965 (F. MacKenzie, AMNH); Cule-
bra, near Dewey, 18°18'N, 65°18’W, 10.viii.1965 (Is-
land Project Staff, Univ. of Puerto Rico, AMNH); Cu-
lebrita Island, 18°19'N, 65°14'W, 15.iv.1965 (H. Hea-
twole & F. MacKenzie, AMNH), 11.viii.1965 (Island
Project Staff, Univ. of Puerto Rico, AMNH); Dese-
cheo Is, 18°23’N, 67°29'W, 19.11.1914 (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.1961 (F. Rolle,
AMNH); Frank Key, Isl. #13 nr. La Parguera area,
17°58'N, 67°3'W, 14.i.1966 (Island Project Staff,
Univ. of Puerto Rico, MCZ); Heatwole Island, off
Culebrita, 18°19’N, 65°13’W, 14.iv.1965 (H. Heatwo-
le & F. MacKenzie, AMNH); Hormiqueros, 18°9'N,
67°S8'’W, 16.11.1962 (Aida Velez, AMNH), 11.iii.1962
(Aida Velez, AMNH), 19.iii.1962 (Aida Velez,
AMNH); Isabela, subexperiment station, 18°30'N,
67°1'W, 17.viii.1957 (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,
(xi LOW (jay Eai€axico,, JEC)LeviniseRockaaial:
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.1958 (A. F.
Archer & M. Sanderson, AMNH); Luquillo, near
beach, 18°23'N, 65°44'W, 24.i1.1932 (A. S. Mills,
AMNH), 24.xii.1985 (V. & B. Roth, CAS), 27.iii.1988
(H. W. & L. Levi, MCZ); Mayagiiez, 18°12'N,
67°9'W, 15.vii.1958 (A. F. Archer, MCZ); Mayagiiez,
5 km N university campus, 18°12'N, 67°9'W, 5.i.1964
(MCZ); Mayagiiez: Las Mesas, 18°11'N, 67°6'W,
20.xi.1960 (F. Rolle, AMNH); Mayagiiez, university
farm N university campus, 18°12’N, 67°9'W, 7.ii.1964
(MCZ); McKenzie Key, Key #3 [P?], 18°12'N,
65°41'W, 29.x.1964 (H. Heatwole, R. Levins & F.
MacKenzie, AMNH); Mona Island, Serrallés, 18°5'N,
67°54'W [P], 7.iv.1944 (Beatty, MCZ); Muertos Is-
land, beating bushes, 17°54'N, 66°32'W, 28.v.1959
(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.1958 (M. W. Sanderson,
AMNH);: Palominitos, 18°20'N, 65°34'W, 16.vi.1965
(Island Project Staff, Univ. of Puerto Rico, AMNH):
Palomino Island, 18°21'N, 65°34'W, 7.xi.1964 (H.
Heatwole & F. MacKenzie, AMNH); Parguera,
17°59'N, 67°3'W, 25.iii.1990 (H. W. & L. Levi, MCZ);
Patillas, 18°0’N, 66°1’W, 3.iv.1931 (Mills & Leonard,
AMNH); Pico Atalaya, Rte. 2, nr. Afiasco, 18°18'N,
67°11'W, 3.vii.1958 (M. W. Sanderson, AMNH); Pi-
fieros Island, 18°15’N, 65°35’'W, 24.ix.1964 (H. Hea-
twole & F. MacKenzie, AMNH); Playa de Humacao,
18°10'N, 65°45'W, 23.v.1964 (MCZ); Quebradillas,
around hotel, 18°29'N, 66°56’W, 30.iii.1989 (H. W. &
L. Levi, MCZ); Ratones Island, 17°56’N, 66°17'W,
19.xi.1964 (H. Heatwole & R. Levins, AMNH); Rom-
ero III [?], 17°57'N, 67°0’W, 13.i.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH); Rubianes,
Caimito Bajo [?], Cord. Jaicoa, 18°26'N, 67°8'W,
19.vii.1958 (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
ponds, 17°56'N, 67°11'W, 23.11.1990 (H. W. & L.
Levi, MCZ); Valle de Lajas, 18°1'N, 67°8'W, 3.vi.1958
(A. F. Archer, AMNH): xeric hills N of Gudnica,
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.1964 (H. Heatwole & F. MacKenzie, AMNH).
U.S. VIRGIN ISLANDS Big Cockroach: 18°24'N,
65°4'W, 7.vi.1966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Congo Cay: 18°22'N, 64°48’W,
12.xi.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Grass Cay: 18°22'N, 64°50'W, 12.xi.1966
(Univ. of Puerto Rico, AMNH). Great Saint James
Island: 18°19'N, 64°50'W, 13.xi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Hans Lollik Is-
land: 18°24'N, 64°55'W, 6.iv.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Little Cock-
roaches: 18°25'N, 65°3’W, 7.vi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Little Hans Lol-
lick: 18°25'N, 64°54'W, 5.iv.1966 (Island Project Staff,
Univ. of Puerto Rico, AMNH). Little Saint James Is-
land: 18°18'N, 64°50'W, 13.xi.1966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Mingo Cay:
18°22'’N, 64°49'W, 12.xi.1966 (Island Project Staff,
Univ. of Puerto Rico, AMNH). Saint John: 18°20'N,
64°45'W, 9.iii. 1925 (F. E. Lutz, AMNH), 11.vii.1958
(A. F. Archer & M. Sanderson, AMNH), 15.xii.1967
(N. L. H. Krauss, AMNH); Cruz Bay, 18°20'N,
64°48'W, 27.11.1964 (A. M. Chickering, MCZ); nr.
Cinnamon & Hart Bays on W half of island, 18°20’N,
64°46'W, 2.viii.1976 (D. E. & D. N. Rosen, AMNH).
Saint Thomas: 18°24'N, 64°55'W, 24.11.1925
(AMNH), 24.vii.1925 (A. Petrunkevitch, CAS),
24.xi1.1925 (AMNH); Crown Mountain, 18°21'N,
64°58'W, 30.viii.1957 (A. F. Archer, AMNH),
7.vii.1958 (A. F. Archer, AMNH); Denmark Hill [?],
18°24'N, 66°55'W, 1.ix.1957 (A. F. Archer, AMNH):
Flagstok Hill, Stumpy Bay, 18°22’N, 65°0'W,
9.vii.1958 (A. F. Archer, AMNH); Harman’s, Char-
lotte Amalia, 18°21'N, 64°56’W, 2.ix.1957 (A. F. Ar
cher & family, AMNH); Hassell Island, 18°20'N,
64°56'W, 1.ix.1957 (A. F. Archer, AMNH), 15.ii.1964
(A. M. Chickering, MCZ), 10.viii.1966 (M. L. Pres-
sick, AMNH). St. Croix: Christiansted, 17°45'N,
64°42'W, 3.vi.191]1 (AMNBH), 15.i1.1955 (A. M. Nad-
ler, AMNH); East End, 17°45'N, 64°40’W, 15.xii.1965
(Island Project Staff, Univ. of Puerto Rico, AMNH);
Green Key, 17°46'N, 64°40’W, 17.iv.1964 (H. Hea-
twole, MCZ); Mount Eagle, dryish forest, 17°46'N,
64°49'W, 15.xii.1965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Protestant Cay, 17°45’N,
64°42’W, 18.iv.1964 (H. Heatwole, MCZ): St. Croix,
17°45'N, 64°54'W, 1.ix.1966 (Chickering, MCZ),
6.ix.1966 (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).
METEPEIRA ® Piel 53
17. Metepeira roraima new species
Figures 141-148, 316; Map 8
Holotype. Female from Rio Surumu, 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
eS,
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. gressa, 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-
er 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.1970 (W. Eber-
hard, MCZ): Cali, around house, 3°27'N, 76°31’W
(W. Eberhard, MCZ); Palmira, 3°32’N, 76°17'W,
1.iii. 1964 (Ballo, CAS); Rio Pance, below Buenos Ai-
res, 3°55'N, 76°8’W, 5.iv.1970 (W. Eberhard, MCZ);
Rio Pance, near Cali., 3°21’N, 76°33’'W, 8.v.1970 (W.
Eberhard, MCZ), 8.vi.1970 (W. Eberhard, MCZ),
15.vi.1970 (W. Eberhard, MCZ), 23.vi.1970 (W.
Eberhard, MCZ), 25.vi.1973 (W. Eberhard, MCZ).
GUYANA Upper Takutu: Isherton, on lat. 2, 10 mi E
Rupununi River, 2°19'N, 59°22’W, 15.xi.1937 (W. G.
Hassler, AMNH).
18. Metepeira gressa (Keyserling)
Figures 149-156, 300; Map 8
Epeira gressa Keyserling, 1892: 166, fig. 123, 2. 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, d.
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-Leitaio, 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), the 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
(Fig. 156). Sternum black with median
white line, 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
1R6:
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. compsa 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
METEPEIRBA ® Piel 55
base and parallel to the embolus tip (Figs.
12932)
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.i.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.ix.1944
(Torres, MLP): Moreno, 34°39'S, 58°48’W, 15.ii. 1966
(Rossi and Maury, MACN); San Isidro, 34°27'S,
58°30'W, 15.xii.1937 (Peregra, MACN); San Mi-
guel—San Fernando, 34°29'S, 58°39'W, 15.vii.1940
(F. Morrios, MACN); Sierra de la Ventana, 38°9'S,
61°48'W, 15.iii.1939 (S. H. Bavio, MACN), 15.xi.1954
(Fritz, MACN), 31.x.1969 (Carlos Grisolia, MCZ),
15.vii.1972 (Amarrilla, MACN); Tigre, 34°25'S,
58°34'W (J. M. Viana, MACN). Entre Rios: Concep-
cién del Uruguay, 32°29'0"S, 58°13'42"W, 4.11941
(Prosen, MLP); Salto Grande, 31°13'S, 57°56’W,
15.iii.1964 (M. E. Galiano, MEG). Santa Fé: Floren-
cia Varelo [?], 28°2’S, 59°15’W, 15.xii.1939 (F. Morris,
MACN). Tucumdén: 30 km S Concepcion, 27°36'S,
65°35'W, 16.1.1983 (L. E. Pehla, AMNH). PARA-
GUAY Itapua: 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.1963 (R. Capocasale and L. Bruno,
CAS). Colonia: Punta Gorda. [?], 34°28'S, 57°51'W,
25.ii.1968 (R. Capocasale and L. Bruno, CAS),
26.ii.1965 (R. Capocasale and L. Bruno, 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. 1983 (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 sharp corner 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 olmec, Metepeira comanche,
Metepeira pimungan, Metepeira triangu-
laris, Metepeira arizonica, Metepeira atas-
cadero, and Metepeira incrassata.
19. Metepeira maya new species
Figures 157-163, 317; Map 14
Holotype. Male from North Bay, Twin Cays, Stann
Creek District, Belize, 14.iii.1986, 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-
teroventral 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 femur
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 forward, 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 597
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, mesal. 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.1986 (P.
Sierwald, USNM); Twin Cays, North part of West
Pond, West of Swamp Doc, 16°50'N, 88°5'W,
28.iii. 1986 (P. Sierwald, USNM); Twin Cays, North-
west Point, North Bay, 16°50'N, 88°5’W, 14.iii.1986
(P. Sierwald, USNM); Twin Cays, Northwest Point,
North Bay, ground not flooded, 16°50'N, 88°5'W,
14.iii.1986 (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.1984 (Feller, USNM), 5.vi.1985 (Erwin,
Mathis, Sims, USNM). COSTA RICA San José: San
José, 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.vili. 1947
(C. & P. Vaurie, AMNH). MEXICO Chiapas: Com-
itan de Dominguez, 16°15'N, 92°8’W, 19.vii.1950 (C.
J. & M. Goodnight, AMNH); near Rio San Gregorio,
between Comitan and Ocotal, 15°45’N, 92°0'W,
18.vii.1950 (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); Tuxtla Gutierrez,
16°45'N, 93°7'W, 10.vi.1964 (Pallister, AMNH). NIC-
ARAGUA Matagalpa: Matagaslpa, 12°53'N, 85°57'W,
15.vii. 1989 (R. Reinbold, JMM).
20. Metepeira inca new species
Figures 164-170, 318; Map 11
Holotype. Male from Parifias 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 ruled Peru.
Description. F emale paratype from Par-
ifias 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 brown 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 medians 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.
maya 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. maya and M. com-
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. comanche
(Fig. 187); the posterior lobes on M. inca
are pointed (Fig. 166), not just thickened,
as in M. maya (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 Piura: 12 mi N Man-
cora, 4°0'S, 80°54'W, 11.xii.1938; Parifias Valley,
4°30'S, 81°8'’W, 3.iv.1939, 8.iv.1939, 16.iv.1939,
7.V.1939, 21.v.1939, 25.vi.1939, 3.vii.1939, 6.viii. 1939,
15.viii. 1939, 15.x.1939; Quebrada de Parifias, 4°32'S,
81°17'W, 14.iv.1939, 7.v.1939, 21.v.1939; Quebrada
Mogollon, 4°32'S, 81°4’W, 30.iv.1939, 11.vi.1939,
18.vi.1939, 21.vi.1939, 11.vii.1939, 16.vii.1939,
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, 2. Female holotype from Pilot Knob Valley,
Mohave Desert, California, in the AMNH, exam-
ined. Roewer, 1942: 868. Bonnet, 1957: 2820. Levi,
1977: 200, figs. 28-36.
Description. Female from Baja Califor-
nia Norte, Mexico. Carapace dirty 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 macrosetae 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
METEPEIRA ° Piel 59
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.1983 (L.
Strange & R. Miller, FSCA). USA Arizona: Show
Low Lake, 34°12'’N, 110°O’W, 15.viii.1966 (MCZ).
22. Metepeira olmec new species
Figures 178-184, 324; Map 11
Holotype. Female from Fortin de las Flores, Vera-
cruz, Mexico, 26.vi.1944, L. I. 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
eS tourtiar 23.
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 spider’s 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 forward, but instead drops off,
forming a sharp curve (Fig. 178). Among
other species in the M. incrassata species
group, this embolus shape is also seen in
M. comanche and M. triangularis: unlike
M. comanche (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 José: San
Antonio de Escanza, 9°59'N, 84°11'W [?], 4.iii.1984
(W. Eberhard, MCZ). MEXICO Veracruz: Fortin 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.1950 (A. M. Chickering,
METEPEIRA ® Piel 61
7
gosoga
comanche
(23)
Figures 171-177. Metepeira gosoga Chamberlin 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 comanche Levi (sp. 23 [185,188,189] 33°22’N, 99°56’W; [186,187] 25°45’/N, 101°55’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.1954 (A. M. Chickering, MCZ). Panama:
Cerro Galera, 8°55'N, 79°38'W, 7.i.1977 (H. W. Levi
& Y. Lubin, MCZ).
23. Metepeira comanche Levi
Figures 185-191, 331; Map 11
Metepeira comanche Levi, 1977: 204, figs. 61-69, ¢,
2. 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 medians 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. maya 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.
maya 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 History. 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
LIEN,
Records Examined. MEXICO Coahuila: Gloria,
26°41'N, 101°23'W, 24.viii.1947 (W. J. Gertsch,
AMNH); Paila, 25°45’N, 101°55’W, 21.viii.1947 (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., Califoria, USA, 20.vii.1968, 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.
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
anteroventral 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 from 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
METEPEIRBA ® Piel 63
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
specimens is curved to form 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 males 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
Coryopsis 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, 11.viii.1995 (W. H.
Piel, MCZ); 34°2'6.1"N, 120°21'9.1"W, 12.viii.1995
(W. H. Piel, MCZ); 34°2’N, 120°22’W, 20.viii.1968
(M. E. Thompson, MCZ); 34°3'3.1"N, 120°21'53.3"W,
13.viii.1995 (W. H. Piel, MCZ): behind dunes on
beach, 34°2'43.8"N, 120°21'0.8"W, 11.viii.1995 (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 labyrinthea:—Franganillo, 1936: 75. Bry-
ant, 1940: 341. Platnick, 1993: 449.
Metepeira triangularis:—Archer, 1958: 15, fig. 37, @.
Metepeira acostai Archer, 1958: 15, fig. 36, 2. Female
holotype from the savannas, Agramont, Camagiiey
Province, Cuba, in the AMNH, examined. NEW
SYNONYMY.
Metepeira bani Archer, 1965: 132, figs. 11, 17 2, 13
3. Male and female syntypes from Bani, Domini-
can Republic, in the AMNH, examined. NEW
SYNONYMY. Brignoli, 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.1954 (J. T. Acosta,
AMNBH).
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
DAD
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 with 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
curve (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.
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 Camagtiey Prov.:
21°30'N, 78°L0’W, x.1954 (J. T. Acosta, AMNH). Ma-
tanzas: Matanzas, Parque Watkins, 23°3'N, 81°35’W,
9.viii.1955 (A. F. Archer, AMNH); Pan de Palenque
[?], 23°1'N, 81°43’W, 11.viii.1955 (A. F. Archer,
AMNH). Oriente: Banes, 20°58'’N, 75°43'W,
2.viii.1955 (A. F. Archer, AMNH); Cuabitas, Santiago,
20°4'N, 75°48'W, 15.xii.1955 (P. Alayo, AMNH):
Puerto Boniato, 20°7'N, 75°47'W, 4.xi.1945 (P. Alayo,
AMNH). Sierra las Casas: Isla de Pinos, 21°53'N,
82°48’W, 17.viii. 1955 (A. F. Archer, AMNH). DO-
MINICAN REPUBLIC Azua: Hatillo, 18°24’'N,
70°32'W, 30.i1.1991 (Felix E. Del Monte, CAS). Ban-
ahona: Sierra Martin Garcia [?], 18°25’N, 70°30'W,
8.viii.1958 (A. F. Archer & E. de Boyrie Moya,
METEPEIRA ° Piel 65
=.
(= 5%)
a 30
4 203
triangularis
(25)
Dios
209 mj 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, 113°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.
206
66 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
AMNRH). La Independencia: 2 km SE EI Limon, Ji-
mani. S side of road, 18°25'’N, 71°45'’W, 30.i1.1991
(Felix E. Del Monte, MNSD); Isla Cabritos: La Des-
cubierta, Lago Enriquillo, 18°34’N, 71°44’W,
18.vi.1981 (E. Marcano, MNSD). Montecristi: Car-
retera Montecristi, El Morro [?], 19°54'N, 71°39'W,
2.xii.1991 (Felix E. Del Monte, MNSD), 21.xii.1991
(Felix E. Del Monte, MNSD). Peravia: Las Dunas,
Prov. Bani [?], 18°25'N, 71°25'W, 23.i.1992 (Felix E.
Del Monte, MNSD). Prov. Trujillo Valdez: W_ of
Bani, 18°17'N, 70°22’W, 8.viii.1958 (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, 2, 3d. Female holotype from Canyon
Lake, Arizona, in the AMNH, examined. Levi,
1977: 200, figs. 12, 13, 39-46, 2, 3. Brignoli, 1983:
OS
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, abdomen, 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 curve (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
Estimated Elevation (meters)
22 24 26 28 30 32 34
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.1973 (S.
C. Williams & K. B. Blair, CAS); 24 mi S Santa Inés,
29°20'N, 114°20'W, 7.vii.1973 (S. C. Williams & K.
B. Blair, CAS); 26 mi S San Felipe, 30°38'N,
114°50’W, 15.iv.1965 (D. Q. Cavagnaro, C. E. & E.
S. Ross, V. L. Vesterby, CAS); 6 km NW Racho Santa
Inés, 29°43'N, 114°43’W, 28.iii.1981 (Paul E. Blom,
WHC); 6 mi S San Felipe, 30°55'N, 114°52’W,
14.iv.1965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V.
L. Vesterby, CAS); 9 km NW Racho Santa Inés,
29°46'N, 114°46’W, 3.vii.1981 (David Crowe, WHC):
Arroyo de Calamajué at carbonated spring, 29°38’N,
114°25'W, 26.vi.1973 (S. C. Williams & K. B. Blair,
CAS); Cafion de Guadalupe, off Laguna Salada,
32°10'N, 115°48’'W, 11.1.1958 (V. Roth, AMNH); El
Mayor, 32°5'N, 115°13’W, 15.vi.1952 (M. Cazier, W.
Gertsch, & R. Schrammel, AMNH); Isla Mejia,
29°34'N, 113°35’W, 30.iv.1921 (J. C. Chamberlin,
MCZ); Isla Partida, 28°53’N, 113°4’W, 2.vii.1921 GE
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 [P], 9.iv.1921 (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.1962 (Howard W. Campbell, CAS);
San Pedro Martir, 30°45’N, 115°13'W, 18.iv.1921 (J.
C. Chamberlin, CAS). Coahuila: 25 mi SE San Pedro,
25°35'N, 102°50'W, 21.viii.1947 (W. J. Gertsch,
AMNH): Cabos, 25°35'N, 101°43’W, 21.viii.1947 (W.
J. Gertsch, AMNH); Gloria, 26°41’N, 101°23’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.1947 (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.1947 (W. J.
Gertsch, AMNH). San Luis Potost: 3 km W. Pilares,
21°55'34"N, 100°48’6"W, 21.x.1994 (W. H. Piel,
MCZ). Sinaloa: Las Saleras Is. (= Isla Saliaca [?]),
25°11'N, 108°20’W [?], 13.vi.1921 (J. C. Chamberlin,
METEPEIRA ® Piel 67
CAS). Sonora: 20 mi N Hermosillo, 29°8’N,
110°58'W, 13.ix.1966 (Jean & Wilton Ivie, AMNH):
20 mi SW Sonoyta, 31°38'N, 113°4'W, 13.vi.1952 (W.
J. Gertsch, AMNH); 22 mi E Hermosillo on the
banks of Rio Sonora, 29°4'’N, 110°55’W, 17.viii.1959
(B. A. Branson, AMNH); Los Angeles, 29°27'N,
110°46'W, 4.x.1966 (V. Roth, AMNH): Puerto Pef-
asco at seashore, 31°20'N, 113°33’W, 3.iv.1968 (D. E.
Bixler, MCZ). Zacatecas: 14 mi S Fresnillo, 23°5’'N,
102°45'W, 4.vili.1954 (W. J. Gertsch, AMNH);: 4 mi
NE Concepcién del Oro, 24°41’N, 101°23'W,
4.vii.1984 (J. B. Woolley, AD). USA Arizona: Santa
Catalina Mtns., mile 7.9 of Catalina Highway from
Tuscon to Mt. Lemmon., 32°17’N, 110°48’W,
19.vi.1985 (W. Maddison, MCZ): Tucson, 32°15’'N,
110°57'W, 5.vii.1991 (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,
B34°11'N, 117°53'W, 19.viii.1964 (L. Pinter, MCZ).
Texas: Big Bend Nat'l Park, Old Ranch House on St.
Elena Rd., 29°10’N, 103°30’W, 25.v.1967 (E. Sabath,
MCZ).
27. Metepeira atascadero new species
Figures 214-221, 336; Map 14
Holotype. 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-
68 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
les founth: 25:
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 scape’s 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 mm.
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 Tascate, 26°12'N,
105°7'W, 27.vii.1947 (W. J. Gertsch, AMNH); Yer-
banfs, 80 mi NW Durango, 24°45'N, 103°50'W,
19.viii. 1947 (W. J. Gertsch, AMNH). Guanajuato: 20
mi E. Guanajuato, 21°1’N, 100°57'W, 15.ix.1976 (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.1985
(MCZ), 26.x.1985 (MCZ). Guerrero: Tecalpulco, 25
km N Iguala, 18°29'N, 99°38’W, 1.1.1948 (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, AMNBH).
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.1967 (R. E.
Leech, REL). Zacatecas: East of Guadalupe,
22°46'N, 102°31'W, 21.viii.1959 (A. F. Archer,
AMNH): Guadalupe, 92°45'N, 102°31'W, 16.viii. 1947
(W. J. Gertsch, AMNH).
28. Metepeira incrassata
F. O. P.-Cambridge
Figures 222-228, 323; Map 9
Epeira Saléi Keyserling, 1864: 93, fig., ¢, 2. Female
holotype from Oaxaca, Mexico, in BMNH, exam-
ined. Keyserling, 1892: 196, fig. 145, 2. Roewer,
1942: 851. NEW SYNONYM Y.
Metepeira incrassata F. O. P.-Cambridge, 1904: 460,
fig. 11, 2. Female holotype from Jalapa, Mexico,
in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2821.
METEPEIRA ® Piel 69
ventura
(29)
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 lIvie (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. 1
Aranea sallei:—F.O.P.-Cambridge, 1904: 519. Roew-
er, 1942: 851.
Araneus incrassata:—FPetrunkevitch, 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 than 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 the
more popular junior name.
Description. Female from Rancho Chu-
la-Vista, Veracruz, Mexico. Brown 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 7.6 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. fount ois.
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. olmec (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. olmec
(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
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.1946 (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.1969 (S. & J. Peck, MCZ). Méx-
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 Judrez, 20°18’N, 97°57'W, 19.iv.1967 (W.
B. Peck, MCZ). San Luis Potost: 10km W Xilitla on
r120, 21°22'N, 99°4'W, 10.viii.1991 (W. H. Piel & G.
S. Bodner, MCZ). Veracruz: 1 mi SW Tlapacoyan,
19°57'N, 97°14’W, 16.vii.1973 (A. Newton, MCZ): 2
mi N Fortin de las Flores, 18°56'N, 97°1'W,
5.viii. 1966 (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.1991 (W.
H. Piel & G. S. Bodner, MCZ): Fortin de las Flores,
18°54’N, 97°O'W, 7.vii.1947 (G. & M. Goodnight,
AMNH), 25.iv.1963 (W. J. Gertsch & W. Ivie,
AMNH), 10.vii.1976 (A. Newton, MCZ), 21.x.1982
(George Uetz, MCZ), 17.vii.1991 (W. H. Piel & G. S.
Bodner, MCZ); Jalapa, 19°32'N, 96°55'W, 14.ii.1948
(H. Wagner, AMNH), 15.iii.1948 (H. Wagner,
AMNH), 15.x.1962 (N. L. H. Krauss, AMNH): Los
Naranjos, 18°21'N, 96°10'W, 4.iii.1948 (H. Wagner,
AMNH): near Monte Blanco, 18°58'’N, 97°1'W,
3.vili.1973 (A. Newton, MCZ): Orizaba, 18°51'N,
97°4'W, 6.vii.1963 (D. Bixler, MCZ); Rancho Chula-
Vista, N of Cérdoba, 19°4'’N, 97°2'W, 18.vii.1991 (W.
H. Piel & G. S. Bodner, MCZ).
METEPEIRA ® Piel ital
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 slim 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 sharp 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 curves 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 k
Figures 229-235, 320; Map 13
Metepeira ensenada Chamberlin and Ivie, 1942: 65,
figs. 166-168, 3. 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, 2. Female holotype, 1 male and 3
female paratypes from between Oxnard and Santa
Monica, California, USA, in the AMNH, examined.
2. 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 median
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 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 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 specimens 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. Norte: 10
m E El Rosario, 30°1L1'N, 115°46’W, 8.vii.1973 (S. C.
Williams & K. B. Blair, CAS); 10 mi N Col. Guerrero,
28°5'N, 114°8’W, 1.ix.1957 (V. Roth, AMNH); 2 mi
SE Eréndira, 31°19’N, 116°19’W, 12.v.1973 (S. C.
Williams & K. B. Blair, CAS); Isla Cedros, Gran Cafi-
on, 28°12'N, 115°15’W, 10.iii.1945 (B. F. Osorio Taf-
all, AMNH): Isla de Cedros, Cerro de Cedros,
28°12'N, 115°15’W, 1.vii.1983 (V. F. Lee, CAS); Isla
de Cedros, Punta Norte [?], 28°22'N, 115°14’W,
3.vii.1983 (V. F. Lee, CAS); Isla de Cedros, trail to
Cerro de Cedros, at spring, 28°12’N, 115°15’W,
27.ix.1984 (D. B. Weissman, V. F. Lee, CAS); Islas
San Benito, Benitos del Oeste, 28°18’N, 115°35’W,
4.vii.1983 (D. C. Lightfoot & V. F. Lee, CAS); Islas
San Benito, Middle Island, 28°19’N, 115°34'W,
9.iv.1981 (Stanley C. Williams, CAS); Islas San Be-
nito, South Island, 28°18’N, 115°35'W, 9.iv.1981
(Stanley C. Williams, CAS); near Consuelo, 6 mi NW
El Rosario, 30°11’N, 115°46’W, 18.iv.1965 (D. Q.
Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS);
Rancho Las Parritas, 10 mi S San Quintin, 30°20'N,
115°57'W, 27.vi.1977 (C. E. Griswold, CAS); Santo
Tomas, 31°33’N, 116°24’W, 8.vii.1953 (W. J. & J. W.
Gertsch, AMNH). Baja Calif. Sur: Bahia de los
Muertos, 23°58'’N, 109°50'W, 20.xii.1958 (H. B.
Leech, CAS); Desierto del Vizcaino, Laguna Ojo de
Liebre, sobre frutilla, 27°43’N, 114°15’W, 11.xi.1981
(A. Cota & M. Jiménez, MLJ); E edge of Sierra Pla-
ceres, 27°35'N, 114°30'W, 25.iii.1984 (W. J. Pulawski,
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.1976 (W. Icenogle,
MCZ); Lompoc, by US 63, 34°38'N, 120°27'W,
17.viii.1966 (L. & P. Pinter, MCZ); Lucia, Monterey
Co., 36°1'N, 121°33'W, 15.ix.1964 (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.1993 (Martin G.
Ramirez & Laura B. Fandino, MCZ):; Santa Catalina
Island, Mt. Torquemada, 33°26’N, 118°33'W,
15.viii. 1965 (L. Pinter, MCZ): Winchester, 33°42'N,
117°5'W, 11.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.1975 (G. F. Knowlton, MCZ).
30. Metepeira revillagigedo new species
Figures 236-242, 301; Map 12
Holotype. Female from south of Isla Socorro, Archi-
piélago de Revillagigedo, State of Colima, Mexico,
15.xii.1988, M. Jiménez, 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, Il. 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-
METEPEIRA ® Piel 73
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
2S:
Male paratype from Isla Socorro, Colli-
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 very 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 very 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.
943).
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 Archipiélago 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 Colima: Archipiéla-
go de Revillagigedo: Isla Socorro, 18°45'N, 110°57’W,
26.iv.1932 (Templeton, Crocker Exped., CAS); Ar-
chipiélago de Revillagigedo: Sur de la Isla Socorro,
18°44'N, 110°57'W, 15.xii.1988 (M. Jiménez, MCZ).
31. Metepeira celestun new species
Figures 243-249, 326; Map 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 with 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 with other species, the
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 from 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-
METEPEIRBA ° Piel 75
Para ry 3 je
ij Pa revillagigedo
239 ¥ 240 (30)
uncata
(32)
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
females 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
IB),
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.1983 (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.1983 (W.
Maddison, MCZ); Balankanche Cave, 2 km E Chi-
chen Itza, 20°40’N, 88°33’W, 19.vii.1983 (W. Mad-
dison, MCZ): beach north of Celesttin, 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.1983 (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 from 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,
fountineoror
Male from Ayutla, San Marcos, Guate-
mala. Carapace dirty yellowish brown with
lighter mark in center (Fig. 253). Same
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
METEPEIRA ® Piel UH
mm. Average body length of four males
examined 5.6 mm, range 4.5 to 6.4 mm.
Natural History. 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.1980 (W.
Eberhard, MCZ). GUATEMALA El Quiche: Chichi-
castenango, 14°56'N, 91°7’W, 6.viii.1947 (C. & P.
Vaurie, AMNH). Huehuetenango: Todos Santos Cu-
chumatan, 15°31’N, 91°37'W, 16.viii.1979 (C. E.
Griswold, CAS). Quetzaltenango: E] Baul, 1 km S
Quetzaltenango, 14°49'N, 91°31'W, 14.viii.1979 (T. C.
Meikle & C. E. Griswold, CAS); Quetzaltenango,
14°50'N, 91°31'W, 16.viii.1950 (C. J. & M. Good-
night, AMNH). San Marcos: Ayutla, 14°40'N, 92°9'W,
19.viii.1947 (AMNH).
33. Metepeira crassipes
Chamberlin and lIvie
Figures 257-263, 309; Map 12
Metepeira josepha Chamberlin and Ivie, 1942: 64, fig.
165, 2. 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, 2, 3. Male holotype, female, male
paratypes from Laguna Beach, California in the
AMNH. Levi, 1977: 202, figs. A= 52 ORO
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,
fOumtle 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 curvature (Fig. 257),
in contrast to much more abrupt curvature
seen in other species in the M. ventura
group (e.g., Fig. 264), with the exception
of M. revillagigedo. 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 History. 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.1953 (W. J.&
J. W. Gertsch, AMNH), 12.xi.1976 (S. C. Williams &
K. B. Blair, CAS). Baja Calif. Sur: 26 mi S Loreto,
95°37'N, 111°17'W, 1.i1.1977 (C. E. Griswold & L.
Vincent, CAS); Isla Magdalena, Puerto Magdalena,
24°38'N, 112°9'W, 16.iii.1957 (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.1921 (G. C. Chamberlin, MCZ); Si-
erra de Alamos, 30°51'N, 112°2'W [?], 19.i.1968 (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.1976 (W.
Icenogle, MCZ).
34. Metepeira chilapae
Chamberlin and lvie
Figures 264-270, 333; Map 13
Metepeira chilapae Chamberlin and Ivie, 1936: 45,
figs. 119-121, d. Male holotype from Chilapa,
Guerrero, Mexico, in the ZMB. Roewer, 1942: 868.
Metepeira chilapica:—Bonnet, 1957: 2820. Unjusti-
fied emendation.
Note. Examination of Chamberlin and_Ivie’s
(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
petatlan
(35)
Figures 257-263. Metepeira crassipes Chamberlin and lvie (sp. 33 [257-261] 33°43’N, 117°6’W; [262,263] 27°58’N, 111°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 chilapae 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 either 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
Dheore
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 anteroventral 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 mm, 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.1984 (J.
B. Woolley, ADC). Morelos: Cocoyoc, 18°52'N,
98°59'W, 28.vii.1956 (W. Gertsch & V. Roth, AMNH);
Cuernavaca, 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.1985 (J. Woolley & G. Zolnerowich,
ADC); Huajuapan, 17°48'N, 97°46’W, 1.x.1946 (Ee
Wagner, AMNH); Oaxaca, 17°3'N, 96°43'W,
17.vii.1955 (C. & P. Vaurie, AMNH), 12.viii.1991 (W.
H. Piel & G. S. Bodner, MCZ); San Balt. Chichica-
pan, 16°45'N, 96°29'W, 4.viii.1991 (W. H. Piel & G.
S. Bodner, MCZ).
Metepeira minima Group
Female spiders in the M. minima group
(Metepeira petatlan, Metepeira minima,
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.
SHTAL. SATS. DASTG,,. OXSIS}).
35. Metepeira petatlan new species
Figures 271-277, 334; Map 10
Holotype. Male from 50 km southeast of Petatlan,
Guerrero, Mexico, 14.viii.1984, J. B. Woolley, in
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 abdomen 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 widening 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 almost 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. Femur 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
METEPEIRBA ° Piel Sl
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 mm, 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. minima 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-
gellum 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.1984 (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,
11.ix.1966 (Jean & Wilton Ivie, AMNH).
36. Metepeira minima Gertsch
Figures 278-285, 328; Map 15
Metepeira minima Gertsch, 1936: 10, fig. 31, ¢. Male
holotype from Edinburg, Texas, in the AMNH, ex-
amined. Roewer, 1942: 869. Chamberlin and Ivie,
1942: 67, fig. 174, 2. Levi, 1977: 206, fig. 70-73,
76-77, 2; 74-75, 6. 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, third 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. minima 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.
DHT Sly)
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
2338
jamaicensis
(38)
&:
294 296 \ 207]
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. HONDURAS Copan: 14°50'N,
89°9'W, 7.ii1.1939 (AMNH): Ruinas, 14°50’N
89°9'W, 7.iii.1939 (AMNH). MEXICO Campeche:
Becan, 18°33'N, 89°30’/W, 31.vii.1991 (W. H. Piel &
G. S. Bodner, MCZ); Campeche, 19°51'N, 90°32’W,
14.vii.1948 (C. J. Goodnight, AMNH); Champotén
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.viii.1949 (C. J. Goodnight, AMNH). Chiapas: 16 mi
W Cintalpa on rt 190, 16°36’N, 93°51’W, 15.vi.1982
(F. Coyle, MCZ). Nuevo Léon: 20 km E Montemo-
relos, Camino Q. Rayones, 25°16’N, 99°41'W,
22.vi.1981 (L. Stange, FSCA): Los Cristales, 25°33’N
100°12'W, 15.viii.1972 (A. F. Archer, AMNH); Villa
de Santiago, Hacienda Vista Hermosa, 25°25'N,
100°9'W, 19.vi.1940 (H. Hoogstraal, MCZ); Villa San-
tiago, Horsetail Falls, 25°25’N, 100°9'W, 19.vi.1940
(H. Hoogstraal, MCZ). San Luis Potost: 20 km W Cd.
Valles, 21°58'N, 99°11’W, 18.iii.1972 (J. A. L. Cooke,
AMNH): Cd. Valles, 21°59’N, 99°1’'W, 6.vii.1940 (P.
Rau, MCZ), 19.vii.1956 (W. J. Gertsch & V. Roth,
AMNH), 15.vii.1959 (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.ix.1956 (A. F. Archer,
AMNH): N section of San Luis Potosi, 22°13’N,
100°58'W, 8.ix.1956 (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°1L0’W,
23.viii. 1965 (W. J. Gertsch & R. Hastings, AMNH).
Tamaulipas: 60 km N Cd Valles, 22°30'N, 99°4'W,
10.viii.1991 (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.1947 (C.
J. & M. Goodnight, AMNH); rl01 26km S Tula,
22°49'N, 99°55’W, 8.ix.1991 (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. ile 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 Helipers 2iles2uNe
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, 9.1.1984
(V. & B. Roth, CAS): Balankanche Cave, 2 km E Chi-
chen Itza, 20°40'N, 88°33’W, 19.vii.1983 (W. Mad-
dison, MCZ); beach north of Celesttin, 20°56’N
90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner,
MCZ); Chichén Itza, 20°40'N, 88°34'W (C. J. Good-
night, AMNH), 15.vii.1981 (C. Gold, CAS): Cordil-
leria Mayapan, 20°28'N, 89°11’W, 6.ix.1952 (J. & D.
Pallister, AMNH); Uxmal, 20°22'N, 89°46’W,
7.ix.1970 (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.1958 (A. Brady,
MCZ); 1 mi S Pharr, 26°10’N, 98°11’W, 14.xi.1958
(A. Brady, MCZ); 1 mi S Pharr on U.S. HW 281,
26°10'N, 98°L1’W, 14.xi.1958 (A. Brady, MCZ).
37. Metepeira pacifica new species
Figures 286-292, 302; Map 10
Holotype. Male from La Pacifica, Guanacaste, Costa
Rica, 1.ii.1975-2.iii.1975, 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 alee 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
Jan
Feb
July I
June
gressa, n = 26
July
June
galatheae, n = 304
Jan
ee
i Feb
i
Sep
July
Apr
“May
June
calamuchita, n = 15
July
June
datona, n = 66
July
June
roraima, n= 11
July June
revillagigedo, n = 2
July June
tarapaca, n = 23
July June
crassipes, n= 8
July June
jamaicensis, n = 24
July June
maya, n= 13
Jan
\ Mar
\
i
|
July
pacifica, n= 15
Oct
Apr
June
Sep
July
XC May
June
rectangula, n= 11
July
June
nigriventris, n = 44
July
glomerabilis, n = 33
Nov al
=~ Jes
Oct IS
June
July
June
inca, n = 29
METEPEIRA ° Piel
85
July
wee"
June
karkii, n = 9
"July
June
compsa, n = 279
July
June
arizonica, n = 45
Figures 300-319. Circular histograms 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 anteroventral 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
eS.
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 Trypar-
gilum nitidum, T. tenoctitlan, and T. ben-
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 Cajfias, La Pacifica, prey of Trypargilum ben-
soni, 10°27'N, 85°9'W, 291.1975 (R. E. Coville,
MCZ); 4 km NW Caijias, La Pacifica, prey of Trypar-
gilum nitidum, 10°27'N, 85°9’W, 1.ii.1975—2.iii.1975
(R. E. Coville, MCZ): 4 km NW Canas, La Pacifica,
prey of Trypargilum tenoctitlan, 10°27'N, 85°9'W,
1.ii.1975 (R. E. Coville, MCZ); 4 km NW Canas, La
Pacifica, wasp collected, 10°27'N, 85°9'W, 16.ii.1975
(R. E. Coville, MCZ); Bagaces, Palo Verde, 10°21’N,
85°21'W, 19.1.1978 (W. Eberhard, MCZ); ca. Cafias,
10°25'N, 85°7'W, 15.xi.1982 (W. Eberhard, MCZ):
Canias, 10°25'N, 85°7'W, 15.xi.1982 (W. Eberhard,
MCZ). Puntarenas: Finca San Miento-Sialas, 10°9’N,
84°54'W, 5.ii.1976 (Roth & Schroepfer, AMNH). San
José [?]: Santa Maria, 9°39'N, 83°58’W, 15.i.1930
(Dodge, MCZ). HONDURAS Tegucigalpa: Teguci-
galpa, 14°6’N, 87°13/W, L.vii.1948 (Clarke, AMNH).
NICARAGUA Managua: Laguna de Jiloa, SW Ma-
nagua, campsite, 12°13'N, 86°19’W, 8.vii.1970 (S.
Riechert, SR).
38. Metepeira jamaicensis Archer
Figures 293-299, 313; Map 15
Metepeira jamaicensis Archer, 1958: 16, fig. 33, °.
Female holotype from Port Henderson, St. Cath-
erine Parish, Jamaica, in the AMNH, examined.
Metepeira minima:—Levi, 1977: 206, 208. Brignoli,
1983: 275. Erroneous synonymy.
Description. F emale from Saint Mary's
Parish, Strawberry Fields near Robin’s Bay
and Green Castle, Jamaica. Carapace dirty
brown, white around eyes, central white
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-
320 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
ventura, n = 35
324 Dec
July
olmec, n= 6
328 Dec
Oct
June
Jan
Feb
Sep
July
minima, n = 39
May
June
July
June
lacandon, n = 10
July
atascadero, n = 17
June
321 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
grandiosa grandiosa, n = 4
July
June
grandiosa alpina, n= 5
July
uncata, n=6
June
333 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
chilapae, n = 15
337 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
triangularis, n = 19
July
gosoga, n= 2
June
July
celestun, n= 8
330 Dec
June
Jan
Sep
July
pimungan, n= 6
June
334 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
petatlan, n = 3
METEPEIRA ® Piel S87
323 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
incrassata, n = 33
327 Dec
July
vigilax, n= 10
June
331 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
comanche, n = 2
335 Dec Jan
Nov Feb
Oct Mar
Sep Apr
Aug May
July June
spinipes, n= 116
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 chilapae;
334, Metepeira petatlan; 335, Metepeira spinipes; 336, Metepeira atascadero; 337, Metepeira triangularis.
88 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
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.1960
(R. A. Lewin, MCZ). HAITI Departement de
L’Ouest: Port-au-Prince, 18°32’N, 72°20'W,
19.vii.1955 (A. F. Archer, AMNBH), 20.vii.1955 (A. F.
Archer, AMNH). Dept. de L’Artibonite: Saint-Marc,
19°7'N, 72°42'W, 15.i.1913 (W. M. Mann, MCZ). JA-
MAICA Cornwall: Montego Bay, 18°28'N, 77°55’'W,
1.iii.1984 (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.1957 (A. M. Chickering, MCZ), 15.vii.1960 (C.
& P. Vaurie, AMNH), 17.vii.1960 (C. & P. Vaurie,
AMNH); Strawberry Fields near Robin’s Bay and
Green Castle, 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.1946
(B. Heineman, AMNH); Saint Ann’s Bay, 18°26'N,
77°8'W, 20.xi.1959 (A. M. Nadler, AMNH). Saint
Catherine: E Green Harbour, S slope of Healthshire,
17°53'N, 76°51'W, 12.viii.l1958 (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,
AMNBH). Surrey: Kingston, Mona Road, pasture,
17°59'N, 76°24'W, 10.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.1960 (C. & P. Vaurie, AMNH). Westmorland:
Negril, 18°16'N, 78°21'W, 24.ii1.1955 (A. M. Nadler,
AMNH): Whitehouse, 18°4'N, 77°58'W, 26.iii.1955
(A. M. Nadler, AMNH).
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INDEX
METEPEIRA ° Piel 91
VIERA, C. 1986. Comportamiento de captura de Me-
tepeira sp. A (Araneae, Araneidae) sobre Acro-
myrmex sp. (Hymenoptera, Formicidae) en con-
diciones experimentales. Arachnologia, 6: 1-8.
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Metepeira sp. A (Araneae, Araneidae). Boletin de
la Sociedsad Zoologica del Uruguay, 5: 5-6.
. 1992. Comparacion de telas de hembras y
juveniles de Metepeira seditiosa (Araneae, Ara-
neidae). Boletin de la Sociedad Zoologica del
Uruguay, 7: 17-18.
VIERA, C., AND F. G. Costa. 1988. Analisis del com-
portamiento de captura de presas por machos
adultos de Metepeira sp. A (Araneae, Araneidae),
utilizando telas de juveniles y hembras adultas
coespecificos. Journal of Arachnology, 16: 141—
152.
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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
Arachnidomyia 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
cancriformis, 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 §, 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
koepckeorum, Araneus 6
92 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
labyrinthea grinnelli, Aranea 34 rectangula, Metepeira 5, 6, 9, 12, 17, 32, 35
labyrinthea grinnelli, Epeira 34 rectangulata, Metepeira 32
labyrinthea grinnelli, Metepeira 34 revillagigedo, Metepeira 16, 19, 73, 75
labyrinthea group, Metepeira 33 roraima, Metepeira 9, 12, 18, 49, 53
labyrinthea, Epeira 21 Saléi, Epeira 68
labyrinthea, Metepeira 7, 9, 10, 14, 17, 32, 34, 46, 48 salei, Metepeira 70
labyrintheus, Araneus 48 sallei, Aranea 70
lacandon, Metepeira 9, 14, 17, 35, 37 sallei, Araneus 70
latigyna, Metepeira 48 santa, Aranea 28
lindae, Arachnidoyia 10 scitulus, Araneus 54
maya, Metepeira 9, 14, 18, 56, 57 seditiosa, Epeira 54
measurements 5 seditiosa, Eustala 54
seditiosa, Metepeira 54
Mecynogea 8, 9
3 seditiosus, Araneus 54
median apophysis §
Metepeira 5 Singa 6
minima group, Metepeira 80 Seer ne et 10 As
minima, Metepeira 9, 10, 12, 16, 19, 81, 83, 86 spinipes, Araneus 34 )
nigriventris group, Metepeira 38 spinipes, Metepeira 5, 6, 8, 9, 14, 17, 34, 35
nigriventris, Araneus 38 suspended remeat © :
nigriventris, Epeira 38 tarapaca, Metepeira 7, 9, 14, 18, 39, 40
nigriventris, Metepeira 9, 10, 11, 12, 18, 38, 39 ae a A wae Shy Ue LS G8), CD
ocosingo, Mecynogea 10 GU ores ;
olmec “Metepeira 9, 16, 19, 59, 61 uncata, Metepeira 9, 16, 19, 75, 76
; : ; uncatus, Araneus 76
i ae ee a ed S304 vaurieorum, Metepeira 48
paomata, Metepenes ventura group, Metepeira 71
palp oy = ventura, Metepeira 10, 16, 19, 69, 71
palustris, Metepeira ao vigilax group, Metepeira 26
perezi, Metepeira 48 vigilax, Araneus 30
petatlan, Metepeira 16, 19, 79, SO vigilax, Epeira 29
pimungan, Metepeira 8, 9, 16, 19, 62, 65 vigilax, Metepeira 9, 10, 12, 17, 26, 30, 31
predation 10 virginensis, Metepeira 48
rayorae, Arachnidomyia 10 web 8
rectangula, Epeira 32 Zygiella 6
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| HARVARD UNIVERSITY VOLUME 157, NUMBER 2
_ CAMBRIDGE, MASSACHUSETTS, U.S.A. 8 JUNE 2001
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
OR DISTRIBUTED BY THE
MUSEUM OF COMPARATIVE ZOOLOGY
HARVARD UNIVERSITY
BrevioraA 1952—
BULLETIN 1863—
Memoirs 1865-1938
Jounsonia, Department of Mollusks, 1941-1974 |
OccCASIONAL 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 ‘llinsietisd 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.
es 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 has 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
INO Stacia oes Ue CE EES ba Ps SAR ee 94
MintnOGuctiOny eats hich ys se Lunes Sibi VU 94
Authors of (ivpesD escripuonss ae 94
Definitions and Organization 96
Taxa Included in This Catalogue __............. 97
ANG SRREN ANC Woy aR Oe he ee 98
@ISINON AG GAGES Se eee 98
Pareallecosy oe SEES Ga 98
mecounts Of Type Specimens 2 99
Order Didelphimeonpiiay se aes 99
HamilysCalucouyidae ees 99
amotly,Maxmiosidact 222.2). eee 99
HannilyaDicdel plidac hase tse = ses eee 100
Order Dasyuromorphia —_......-.-.---------------- 100
[Seilh\y IDE tials: A 100
OxdemPerame lia ee ee 100
Hamnilyaeeronyctidac sss ka sleelee olla 100
Ordem@ingulatae ee sais wearer 101
HamulyDasypodidacs a ee 101
OrdemAtnosoncida == = ame ee 101
Hamnily Chrysochlonidae == a. - 101
Hannilyem@emkecidae 2:0 ee 101
Orders Rodentiay= ==. ee ee 103
HamiulyeAplodontidacy== 2s" 2 ssnurs ls 103
HamulysSciumGac, a. sai e te Be 103
Hamulya@astondac ge 2. Bees 109
anulvaGcommidac. =. sere) eee 110
annilvalteteromiyidae sess eee ee 111
lSeneotllhy ID Mhovorettoleyss Was 112
I Semanullhy, Ilona es: eee Ea)
Sulotfannilysarvicolimae sus eenen nee ILD
Sulbtaunilyz@nicetinac =e ees LIE
Subfamily Cricetomyinae ____..-_- LAL
Subfamily Dendromurinae —__-- LAL
Subfamulya@erbillinae = ILE 7e
SWlonehoanllhys IY Munabieves Se 117
Subtamuily Nesomyimae 9 == 120
Subtamulya@ tomy aes eee ee 120
Subfamily Sigmodontinae _....._....-.-...- 121
Hamnilyakedetidac)s ==) eee 128
‘Mammal Department, Museum of Comparative
Zoology, Harvard University, Cambridge, Massachu-
setts 02138.
HamnilySNiyOxiG. ae ee eeeeete sae En mL anenun me
HamulysBathyercidac =u ln stm aes
amnilyeHraithiZomtiGdacr re ear eee
HamilysDasyproctidac: 2 = a ane
HamnilyeAcoutiGaey= sane fae es ae rneE ee
Harnily; Octodomtidacise= set ee
arailyais chimayi dace. eels eee n a
HamnilysCapnomnyi dae jes ee
Orders Bagomompnay se ee ee
Mamnilys@chotomidae = == snsss men
Benrmllyy JUsteroa CHS) oe ee
Order Scandentiay ee
arnilyaslupaidacy ues asses eaeeeneen
Ovrcler IPatmbiveg a
Hammill dae gas oe, oe eee en ee careers
HamuilysDaubentoniidac) 9. sss os
HannilysG alagonidac sass: eee eee
Hamnilya@ebidac geass eo ee eee:
Hamulyaetylobatidac ttm. semen ie ee
EamnilyarlomuniGd acy 22s salsa ine eee
@rderplaipotyplila ss8 os eae eee
Hamily)Nesophontidac (22. ts ee
Family Solenodontidae 02.2228
[stangaullyy: SYO;atGNG Vs) Ue ee al
tenooulby IRalhonGhys)) es eee
Order Chiropterass -iiey i ees
amilyaPteropodidae) = aasaes .. slr aasnmens
anuly SH mlbjallonuridac) parents. ero
Jenene hyowerate evenee eS a
Hamilynninolophidac === ==
HannilyaNormnoopidac === s=aesananenenas
Hamulysehyllostomidac === ===
HamailygMolossiclac yams see sean
HamulyaVespenrtilionidae) =a: sass aa
mewaavilye TU oyjanoyoueratolevey Se et
Order Antiodac ty ats. ss eee nee
Family Tayassuidae ESR ORY Ae EAE ES
Hamuilya Monod ontidac yas es seen
Hannilyathocoemicd ac eee maen aaleene
Iaeioalhy CRIA: a
IRlemodtly” JEN ANCEYS ie es
Orrgler Carnivora) se
Banari (CRNA YS) op
HanailyaWinsidac heen senate: seen eece sees
Bull. Mus. Comp. Zool., 157(2): 93-181, June, 2001
94 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family Procyonidae PSS late Aare Nc nate TO 151
HamulvaMiustelicacmems a: sa ennasenenes 152
|@eronvilhy: IN iejo) owt ORS 154
| Bertani hyn WaKeranKC ENS) Qo 155
Mamaily Eero eS ti ac aes eee 155
Benoa iy JSC RS =a 155
OxdemEinolestaya eee 156
HanailyaNiani ac aes ass see ee eee ane 156
RE TENE TNC Sp eisie a tas tees a cect ce an ORE COLE IN 157
IIFas Vey: co poh Bee SN Wf eer ets ties etal Tae Toes cha 171
ABSTRACT. The Mammal Department at the Mu-
seum of Comparative Zoology houses name-bearing
types of 342 species-group taxa of Recent mammals.
The type collection consists of 327 holotypes, 2 lec-
totypes, 4 complete syntype series, and 9 partial syn-
type series. This catalogue notes infor mation on the
type locality, collector, dae 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 gteat mammalogist
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 mammals 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 muse-
um’s holdings of mammal type specimens
have grown considerably. Furthermore,
type specimens of mammals 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-
tematic investigations; accordingly, “every
institution in which name-bearing types
are deposited should publish lists Be name-
bearing types in its possession or custody”
(International Commission on Zoological
Nomenclature, 1999: 79). Our duty in this
regard is thus long overdue, and this cat-
alogue should serve to fulfill that obliga-
tion. Without doubt, it will alleviate con-
fusion in the scientific community as to the
whereabouts of certain specimens, thought
to be lost, especially those type specimens
that were acquired from Guillaume Gran-
didier. In addition, it should bring to light
information, formerly unavailable, on this
museum's very notable holdings of name-
bearing types.
All taxonomic 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 mammalogists 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 published 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 legendary, 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 Bonn
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 whales 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 History
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 History, 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 mammal 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 many 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-
ng types.
Locality
This category 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 mammals in the American Mu-
seum of Natural History (Lawrence,
1993). For systematic consistency, the tax-
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden Of
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, Lagomorpha,
Scandentia, Primates, Lipotyphla, Chirop-
tera, Artiodactyla, Carnivora, and Cimoles-
ta. This sequence represents the ongoing
understanding of higher mammalian 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 unnecessary 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 history
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 deserves some mention. When it was
received by the Mammal Department, Guil-
laume Grandidier’s 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 within this catalogue in separate
accounts. The remaining specimen, a fruit
bat, MCZ 45073, is marked exactly like the
other type specimens, and both its skin and
its skull tags bear the name “Eidolon saka-
lava nov. spec., G. Grandidier” in the script-
ed handwriting typical of Grandidier’s 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 this taxon or any ref-
erence to this name in the literature. This
name is therefore invalid, barring a discov-
ery in the future that it was indeed pub-
lished and has since been overlooked. What-
ever the published status of the name, it ap-
pears to me that this specimen should be
considered as a young specimen of Eidolon
dupreanum rather than a distinct species,
and discussion of this mysterious binomial
here should in no way be construed 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
History, New York
BMNH_ Natural History Museum, Lon-
don
BSNH — Boston Society of Natural His-
tory, Boston
FMNH_ Field Museum of Natural His-
tory, Chicago
MNHN Muséum National d Histoire
Naturelle, Paris
MVZ Museum of Vertebrate Zoology,
Berkeley, California
RMNH_ Rijksmuseum van Natuurlijke
Historie, Leiden
USNM_ National Museum of Natural
History, Washington, D.C.
YPM 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-
ner, Judith Chupasko, Jane Harrison, and
especially Maria Rutzmoser for access to
the type specimens. Mary Sears, Timothy
McNeece, and Ronnie Broadfoot at the
Ernst Mayr Library, Harvard University,
were extremely helpful in pinning down
difficult references. We also thank Alison
Pirie for allowing us to make use of the
MCZ Ornithology Department locality
references; Carolyn Kirdahy of the Boston
Museum of Science, who 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 =Neotoma fuscipes fus-
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-
sularis 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 ¢ Helgen 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.
Sciurus castanotus Baird, 1855 and Sciurus castan-
onotus Baird, 1858 =Sciurus aberti aberti 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 (Sciwrus castanotus Baird,
1855 and Sciuwrus castanonotus Baird, 1858), for
taxonomic consistency. MCZ 4692 is a paralecto-
type.
Spermophilus obsoletus Kennicott, 1863 =Spermo-
philus spilosoma obsoletus Kennicott, 1863. Lec-
totype, 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.
Spermophilus parryi var. kodiacensis J. A. Allen, 1874
=Spermophilus parryi 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.
Spermophilus tridecemlineatus var. pallidus J. A. Al-
len, 1874 =Spermophilus 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 list 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 Cary
(1906: 88). G. M. Allen (1931: 255) provided a list
of paralectotypes.
ACCOUNTS OF NAME-BEARING TYPE
SPECIMENS
Order DIDELPHIMORPHIA Gill, 1872
Family CALUROMYIDAE Kirsch and Reig,
1977
Genus CALUROMYS J. A. Allen, 1900
Philander cicur Bangs, 1898k
Proc. Biol. Soc. Washington, 12: 161, 10
August.
=Caluromys lanatus cicur (Bangs, 1898).
See Cabrera (1958: 2).
Holotype. BS8114. 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.
Type 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, 1898i
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:
DIL),
Marmosa mitis Bangs, 1898k
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.
Type 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 (BS8118 is at Wellesley College, B8124
and B8141 are at FMNH, and B8136 and B8138
are at USNM).
100
Marmosa fulviventer Bangs, 1901b
Amer. Nat., 35: 632, 22 August.
=Marmosa robinsoni fulviventer Bangs,
1901. See Handley (1966: 775).
Holotype. B8435. Skin 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. Skin 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/IDELPHIS 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, 1898b
Proc: Boston’ Soc. Nat. Hist., 28: 172, 15
March.
Holotype. B3500. Skin and skull. Adult female.
Locality. (United States): Florida, Brevard County,
Oak Lodge, East Peninsula opposite Micco. 31 Jan-
uary 1895.
Collector. O. Bangs.
Condition. Skin and skull complete.
Type Series. 11 paratypes; all represented by skin
and skull, most still in the MCZ.
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 DASYUROMORPHIA Gill, 1872
Family DASYURIDAE Goldfuss, 1820
Genus ANTECHINUS Macleay, 1841
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Antechinus mayeri misim Tate, 1947
Bull. Amer. Mus. Nat. Hist., 88: 130, 20
February.
=Antechinus naso misim Tate, 1947. See
Laurie and Hill (1954: 7).
Holotype. MCZ 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; MCZ 29923, skin 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 Flannery (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 MYOICTIS Gray, 1858
Myoictis melas wavicus Tate, 1947
Bull. Amer. Mus. Nat. Hist., 88: 140, 20
February.
=Myoictis melas wallacei Gray, 1858. See
Flannery (1990: 56).
Holotype. MCZ 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. Skin and skull complete.
Type Series. Holotype 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. MCZ 7006. Skin and skull. Adult.
TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden
Locality. (Indonesia): Dutch New Guinea (=Irian
Jaya), Doreh Bay, Manokwari, “not far from the
foot of Mt. Arfak.” 23 February 1907.
Collector. T. Barbour.
Condition. Skin complete. Skull partial (occipital
region and two posterior upper molars on each side
missing).
Type Series. Holotype only.
Comments. S. hispida is the type species of Swil-
lomeles G. M. Allen and Barbour, 1909.
Echymipera rufescens australis 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 McIlwraith Ranges, Rocky Riv-
er, “Rocky Scrub.” 20 June 1932.
Collector. P. J. Darlington, Jr., Harvard 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 Flannery (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 Illiger, 1911
Family DASYPODIDAE Gray, 1821
Genus DASYPUS Linnaeus, 1758
Dasypus novemcinctus hoplites G. M.
Allen, 1911a
Bull. Mus. Comp. Zool., 54: 195, July.
Holotype. MCZ 8116. Skin, skull, and_posteranial
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).
101
Order AFROSORICIDA Stanhope et al.,
1998
Family CHRYSOCHLORIDAE Gray, 1825
Genus CHRYSOCHLORIS Lacépéde,
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-
guru 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. Holotype only.
Comments. Included in Chrysochloris stwhlmanni
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
January.
= Geogale aurita Milne-Edwards and A.
Grandidier, 1872. See Genest and Petter
GiOi5:23))
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 § 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 Cryptogale 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-
nerive (=Fenoarivo Atsinanana). April 1928.
Collector. R. Decary. Original number 12.
Condition. Alcoholic, cranium complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Ge-
nest and Petter (1975: 3). The holotype of G. a.
orientalis seems to be the only record of Geogale
aurita from the east coast of Madagascar.
Genus MICROGALE 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 Paléontologie,
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)
synonymized drouhardi with Microgale cowani, an
approach followed by Hutterer (1993: 71), but Jen-
kins et al. (1997: 6) argued that M. drouhardi 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.
Type 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 prolixacaudata 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.
Paramicrogale occidentalis G. Grandidier
and Petit, 1931
Bull. Soc. Zool. France, 56: 129, 15 June.
=Microgale brevicaudata 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 Froriep, 1806
Dasogale fontoynonti G. Grandidier,
1930a
Bull. Acad. Malgache, n. ser., 11: 85 (for
1928).
= Setifer setosus (Schreber, 1777). See
Poduschka and Poduschka (1982: 261).
Holotype. MCZ 45016. Skin, skull, and posteranial
skeleton. Juvenile.
Locality. Madagascar: east coast. 1917.
Collector. Received by G. Grandidier from the
Académie Malgache in 1917.
Condition. Skin represented by a small patch of
fur. Skull partial (occipital region missing). Postcra-
nial skeleton complete, partially articulated.
Type 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
(1987b: 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. Skin and skull. Adult male.
Locality. (Canada): British Columbia, Hope,
Roab’s 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 medialis 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 lachrymal_ bro-
ken—bulla present). Mandible disarticulated.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Med-
way (1977: 90).
Callosciurus ferrugineus 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 (45S m). 20
February 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,
skull, adult male.
skin and
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 February 1911.
Collector. Kobayashi Collection. Original number
AG.
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 Zoology, Vol. 157, No. 2
Genus DREMOMYS Heude, 1898
Dremomys pernyi flavior G. M. Allen,
1912c
Proc. Biol. Soc. Washington, 25: 178, 24
December.
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).
Holotype. 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. Holotype only.
Sciuropterus alpinus lascivus Bangs,
1899}
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. W. Price and P. O. Simons. Original
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.
number 1722.
Condition. Skin and skull complete.
Type Series. 2 paratypes, B9187, skin and skull, fe-
male; B9188, skin and skull, female.
Comments. G. s. lascivus was retained as a valid
subspecies by Hall (1981: 451).
Locality. China: Hupeh (=Hubei), Ichanghsien,
Nantou. 5 February 1909.
Collector. W. R. Zappey. Original number 373.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 7583, skin and skull,
adult male.
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
Genus FUN/ISCIURUS Trouessart, 1880
Funisciurus pyrrhopus victoriae G. M.
Allen and Loveridge, 1942
Bull. Mus. Comp. Zool., 8: 180,
February.
=Funisciurus pyrrhopus akka 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 GLAUCOMYS 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. Skin and skull. Adult male.
Locality. (United States): Idaho, Idaho County.
March 8, 1897.
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. s. 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. Skin and skull. Adult male.
Locality. (United States): West Virginia, Greenbri-
TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden
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, 1896}
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. Skin 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, 1908
Pteromys (Hylopetes) alboniger orinus G.
M. Allen, 1940
The Mammals of China and Mongolia,
Natural History 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 Coolidge, 1940
Bull. Mus. Comp. Zool., 87: 153, 31
December.
=Hylopetes phayrei anchises (G. M. Allen
and Coolidge, 1940). See Ellerman and
Morrison-Scott (1951: 469).
Holotype. MCZ 35776. Skin and skull. Adult male.
Locality. Siam (=Thailand): (Chiang Mai) Mount
105
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 skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 35775, skin and
skull, 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 skull 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
Collector. W. W. Brown, Jr. Original number 631.
Condition. Skin and skull complete.
Type Series. 4 paratypes; MCZ 10405, adult fe-
male, 10406, adult female, 10407, adult female,
10408, juvenile male; all represented by skin and
skull.
Comments. M. a. browni was retained as a valid
subspecies by Hall (1981: 439).
Genus PARAXERUS Forsyth Major, 1893
Aethosciurus byatti laetus G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. Zool., 75: 96,
February.
= 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 February 1930.
Collector. A. Loveridge.
Condition. Skin 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 Miller, 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. Skin 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,
the skull of the type was lost.” The skin and skull
have been reunited subsequently.
Genus SC/URUS Linnaeus, 1758
Sciurus (Guerlinguetus) aestuans
chiriquensis 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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. Soc. Washington, 10: 158, 28
December.
Holotype. B4519. Skin 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 Koprowski (1994: 1).
Sciurus carolinensis var. yucatanensis J.
A. Allen, 1877
In Coues and J. A. Allen, Monogr. N.
Amer. Rodentia, U.S. Geol. Geograph.
Survey 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. Skin 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, 1896)
Proc. Biol. Soc. Washington, 10: 150, 28
December.
= Sciurus niger vulpinus Gmelin, 1896.
See Barkalow (1954: 25).
Holotype. B5215. Skin and skull. Adult female.
Locality. (United States): West Virginia, Greenbri-
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
er County, White Sulphur Springs. 29 January
1896.
Collector. T. Surber. Original number 55.
Condition. Skin 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 Hershkovitz (1947: 37).
Holotype. 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 skin.
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, 1900d
Proc. New England Zool. Club, 2: 43, 20
September.
= Sciurus granatensis morulus Bangs,
1900. See Miller and Kellogg (1955:
DT):
Holotype. B8420. Skin and skull. Adult female.
Locality. Panama: Canal Zone, Loma del Leon. 13
March 1900.
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. morulus was retained as a valid
subspecies by Hall (1981: 437) and Nitikman
(1985: 1).
Sciurus variabilis saltuensis Bangs, 18980
Proc. Biol. Soc. Washington, 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. Skin and skull complete.
Type Series. 2 paratypes; B8145, skin and skull,
adult male; B8244, skin and skull, adult female.
Comments. S. g. saltwensis was retained as a valid
107
subspecies by Cabrera (1961: 368) and Nitikman
(1985: 1).
Genus SPERMOPHILUS F. Cuvier, 1825
Citellus obscurus siccus G. M. Allen, 1925
Amer. Mus. 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 (=Shenxi), 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.
Type Series. Holotype only.
Spermophilus armatus Kennicott, 1863
Proc. Acad. Nat. Sci. Philadelphia, 15:
158, June.
Syntypes. 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
description; 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 armatus 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 the number USNM
3472, is not to be found in the collections of either
museum.
108
Spermophilus elegans Kennicott, 1863
Proc. Acad. Nat. Sci. Philadelphia, 15:
158, June.
Syntypes. 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. Drexler.
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 of 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 Spermophilus armatus. S. elegans is con-
sidered a valid species by Hoffmann et al. (1993:
446) and Nowak (1999: 1254).
Spermophilus (Ictidomys) tridecemlineatus
badius Bangs, 1899c
Proc. New England Zool. Club, 1: 1, 8
February.
= 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-
ymized badius with texensis under the genus Ci-
tellus.
Genus SYNTHEOSCIURUS Bangs, 1902
Syntheosciurus brochus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 25, April.
Holotype. MCZ 10402. Skin 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Condition. Skin complete. Skull partial (left supra-
occipital damaged)
Type Series. 1 paratype; MCZ 10403, skin and
skull, adult female.
Comments. Type species of the genus Syntheosciu-
rus 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 TAMIAS llliger, 1811
Tamias cooper! 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. Skin 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 explained 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 January 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 Mexico (Grant
County), “Fort Webster, Coppermines of the 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 * Helgen and McFadden
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. Skin and cranium.
Locality. (Canada): (Ontario), eastern end of Lake
Superior (near mouth of Montreal River). 5 July
1848.
Collector. L. Agassiz.
Condition. Skin 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, 1896h
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 missing).
Type Series. 2 paratypes; B5479, skin and _ skull,
adult female; B5605, skin and skull, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 340).
Genus TAMIASCIURUS Trouessart, 1880
Sciurus hudsonicus gymnicus Bangs,
1899f
Proc. New England Zool. Club, 1: 28, 31
March.
= Tamiasciurus hudsonicus gymnicus
(Bangs, 1899). See Osgood (1938: 438).
Holotype. B4914. Skin and skull. Adult female.
Locality. (United States): Maine, Piscataquis
109
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, 1896}
Proc. Biol. Soc. Washington, 10: 161, 28
December.
= Tamiasciurus hudsonicus loquax
(Bangs, 1896). See A. H. Howell (1936:
Ih)
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, 1897i
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 skin and skull, 9 females
and 4 males.
Family CASTORIDAE Hemprich, 1820
Genus CASTOR Linnaeus, 1758
Castor caecator Bangs, 1913
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
Type Series. Holotype only.
Comments. C. c. caecator was retained as a valid
subspecies by Hall (1981: 602).
Family GEOMYIDAE Bonaparte, 1845
Genus GEOMYS 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),
Holotype. B5001. Skin and skull. Adult male.
Locality. (United States): Georgia, Camden Coun-
ty, Arnot Plantation, about 4 miles (6.4 km) west
of St. Mary’s. 21 March 1896.
Collector. O. Bangs. Original number 8.
Condition. Skin 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 Williams 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. Skin 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
fit B5016, not B5015. B5016 has “Type” written
after the entry 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 cumberlandius.
Geomys floridanus austrinus Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 177, 15
March.
=Geomys pinetis pinetis Rafinesque,
1817. See Williams and Genoways (1980:
444).
Holotype. B6983. Skin and skull. Adult male.
Locality. (United States): Florida, Pinellas County,
Belleair. 3 August 1897.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Collector. W. S. Dickinson.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype, in the USNM.
Geomys tuza goffi Sherman, 1944
Proc. New England Zool. Club, 23: 38,
30 August.
=Geomys pinetis goffi Sherman, 1944.
See Harper (1952: 37).
Holotype. B7222. Skin and skull. Adult male.
Locality. (United States): Florida, Brevard County,
Eau Gallie. 1S March 1897.
Collector. O. Bangs. Original number 1.
Condition. Skin and skull complete.
Type Series. 12 paratypes; B7212—B7217, B7219—
B7221; all represented by skin and skull, 5 females
and 7 males.
Comments. Williams and Genoways (1980: 444)
synonymized gofft with G. pinetis pinetis; however,
Hall (1981: 504) maintained G. p. goffi as a valid
subspecies. The IUCN designates goffi as extinct.
Genus ORTHOGEOMYS Merriam, 1895
Macrogeomys cavator Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 42, April.
= Orthogeomys cavator cavator (Bangs,
1902). See Russell (1968a: 532).
Holotype. MCZ 10381. Skin and skull. 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-
articulated.
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.
= Orthogeomys 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 * Helgen and McFadden
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 29035, skin and
skull, juvenile female; MCZ 29039, skin and skull,
unsexed juvenile.
Comments. Retained as a valid subspecies by Hall
(1981: 509).
Genus PAPPOGEOMYS Merriam, 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 skull. Adult male.
Locality. Mexico: San Luis Potosi, near San Luis
Potosf, Soledad, 6,400 ft (1,952 m). 1 August 1923.
Collector. W. W. Brown, Jr. Original number 231.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large paratype series in the
MCZ.
Comments. P. c. rubellus was retained as a valid
subspecies by Hall (1981: 520).
Genus THOMOMYS Wied-Neuwied, 1839
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).
JEILIL
Family HETEROMYIDAE Gray, 1868
Genus DIPODOMYS Gray, 1841
Dipodomys californicus pallidulus Bangs,
1899)
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, skin and skull,
adult female.
Dipodops ordii palmeri J. A. Allen, 1891
Bull. Amer. Mus. Nat. Hist., 3: 276, 30
June.
=Dipodomys ordii palmeri (J. 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. EF. Palmer.
Condition. MCZ 5886: Skin 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. 0. palmeri was retained as a
valid subspecies by Hall (1981: 569).
Genus HETEROMYS Desmarest, 1817
Heteromys repens Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 45, April.
=Heteromys desmarestianus repens
Bangs, 1902. See Goldman (1920: 115).
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. Skin 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
Genus LIOMYS Merriam, 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 February 1878.
Collector. E. Palmer.
Condition. Skin complete, skull within skin.
Type Series. Holotype only.
Comments. L. i. alleni was retained as a valid sub-
species by Hall (1981: 590).
Genus PEROGNATHUS Wied-Neuwied,
1839
Perognathus longimembris bangsi 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. Skin complete. Skull partial (teeth sep-
arate from skull). Mandible disarticulated.
Type Series. 2 paratypes; B5302, adult male, skin
and skull; 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 hardyi Batchelder, 1899
Proc. New England Zool. Club, 1: 5, 8
February. :
=Zapus hudsonicus acadicus (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.
Type Series. There is a series of paratypes in the
MCZ.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Zapus hudsonius ladas Bangs, 1899d
Proc. New England Zool. Club, 1: 10, 28
February.
Holotype. B4169. Skin and skull. Adult female.
Locality. (Canada): Labrador, Hamilton Inlet, Ri-
goulette. 15 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 orarius Preble, 1899
N. Amer. Fauna, 15: 29, 8 August.
=Zapus trinotatus orarius 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: $46) and Gannon (1988: 1).
Family MURIDAE Illiger, 1811
Subfamily ARVICOLINAE Gray, 1821
Genus CHIONOMYS Miller, 1908
Hypudaeus nivicola Schinz, 1845
Syst. Verzeichniss Saiugethiere 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. Skin complete.
Type Series. Schinz wrote that he had examined 16
specimens, “junge und alte.”
Comments. G. M. Allen discussed the somewhat
uncertain type status of this specimen (1931: 263).
The original label bears the following: “Hypodaeus
[sic] nivicola juv. 2 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 LES
Genus CLETHRIONOMYS Tilesius, 1850
Evotomys proteus Bangs, 1897g
In Bailey, Proc. Biol. Soc. Washington,
11: 137, 13 May.
=Clethrionomys gapperi proteus (Bangs,
1897). See Jackson (1938: 433).
Holotype. B4081. Skin 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
the 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 exsul G. M. Allen, 1919a
Bull. Mus. Comp. Zool., 62: 532,
February.
Holotype. MCZ 11885. Skin and skull. Adult male.
Locality. (United States): (Alaska), St. Lawrence 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 Nowak (1999: 1479).
Genus EOTHENOMYS Miller, 1896
Craseomys aquilus G. M. Allen, 1912b
Mem. Mus. Comp. Zool., 40: 216,
August.
=Eothenomys 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. Skin complete. Skull partial (hole in left
parietal, occiput missing).
Type Series. 5 paratypes; MCZ 7189, 7191-7194,
7196; all represented by skin and skull, all female.
Microtus (Eothenomys) aurora G. M.
Allen, 1912b
Mem. Mus. Comp. Zool., 40: 211,
August.
=Eothenomys melanogaster aurora (G. M.
Allen, 1912). See Hinton (1923: 149).
Holotype. MCZ 7788. Skin and skull. Male.
Locality. China: Hupeh (=Hubei), Changyangh-
sieh. 2 February 1909.
Collector. W. R. Zappey. Original number 372.
Condition. Skin and skull complete.
Type Series. 3 paratypes; MCZ 7185, skin and skull,
female; MCZ 7186, skin and skull, male; MCZ
7188, skin and skull, male.
Comments. E. m. aurora was retained as a valid
subspecies by Zhang et al. (1997: 229).
Microtus (Eothenomys) mucronatus G. M.
Allen, 1912b
Mem. Mus. Comp. Zool., 40: 214,
August.
=Eothenomys melanogaster melanogaster
(Milne-Edwards, 1871). See G. M. Allen
(1940: 806).
Holotype. MCZ 7789. Skin and skull. Adult 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, skin and skull,
adult female (exchanged to the BMNH); MCZ
7791, skin and skull, adult female; MCZ 7803, skin
and skull, juvenile female.
114
Genus LEMMUS Link, 1795
Lemmus paulus G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 60, 9
April.
=Lemmus sibiricus chrysogaster J. A.
Allen, 1903. See Ellerman and Morrison-
Scott (1951: 656).
Holotype. MCZ 15268. Skin and skull. Adult male.
Locality. (Russian Federation): northeastern Sibe-
ria, Kalaschowo, near mouth of Kolyma River. 22
June 1912.
Collector. J. Koren. Original number 152.
Condition. Skin complete. Skull partial (left man-
dibular ramus missing).
Type Series. 1 paratype; MCZ 15267, skin, female.
Genus MICROTUS Schrank, 1798
Arvicola breweri Baird, 1858
Mammals, in Repts. Explor. Surveys
Railr. to Pacific, 8(1): 525, 14 July.
=WMicrotus breweri (Baird, 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 syntype of breweri, could not
be found.
Arvicola riparia var. longipilis Baird, 1858
Mammals, in Repts. Explor. Surveys
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. Skin complete. Skull partial (part of pa-
rietal, both tympanic bullae and palatine missing).
Mandible disarticulated.
Type Series. See comments.
Comments. G. M. Allen (1931: 260), mentioned
that “although no individuals are mentioned [in
Baird’s description], ... [this] specimen was un-
doubtedly among those examined by Baird in pre-
paring his diagnosis and, therefore, is a cotype
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
[=syntype].” Baird’s specimens were collected by
Kennicott at West Northfield, [linois, 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 syntypes.
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. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. M. p. terraenovae was retained as a val-
id subspecies by 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 complete. Mandible dis-
articulated.
Type Series. Paratype material consists of Bangs’
series from the type locality; B7952—7967, all rep-
resented by skin and skull, 10 females, 3 males, and
3 unsexed individuals. B7953 and B7955 were 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 * Helgen and McFadden
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 paratypes in
the MCZ.
Comments. M. p. enixus was retained as a valid
subspecies by Hall (1981: 793).
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.
Locality. (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.
Type Series. 7 paratypes; B3838—B3544; all repre-
sented by skin and skull, 2 females and 5 males.
Comments. M. p. fontigenus was retained as a valid
subspecies by Hall (1981: 793).
Microtus koreni 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] shattucki
Howe, 1901
iRnoe Portland Soc) Nat. Iaist.. 2 2012 31
December.
=Microtus pennsylvanicus shattucki
Howe, 1901. See Wyman (1922: 162).
Holotype. MCZ 10011. Skin and skull. Adult female.
Locality. (United States): Maine, Penobscot Bay,
115
near Long Island, Tumble 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, Digby. 22 July
1894.
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: 792).
Microtus provectus Bangs, 1908
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. Skin 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 MYOPUS Miller, 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. Skin and skull. Adult male.
Locality. (Russian Federation): northeastern Sibe-
ria, Yakutsk, Nijni Kolymsk (=Nizhnekolymsk),
near mouth of Kolyma River. 28 March 1912.
Collector. J. Koren. Original number 264.
Condition. Skin and skull complete. Mandible dis-
articulated.
116
Comments. 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 zibethicus 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, skin and skull,
adult female; B7975, skin 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
Proce, Bostom Soc. Nat Hist) 26:54" 31
July.
= Ondatra zibethicus rivalicius (Bangs,
1895). See Davis and Lowery (1940:
DAD)
Holotype. B2719. Skin and skull. Adult male.
Locality. (Canada): Louisiana, Plaquemines Parish,
Burbridge. 31 January 1895.
Collector. F. L. Small. Original number 1556/165.
Condition. Skin 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, skin and skull,
male.
Comments. O. z. rivalicius was retained as a valid
subspecies by Hall (1981: 827).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus PHENACOMYS Merriam, 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. Skin 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 Baird, 1858
Synaptomys fatuus Bangs, 1896d
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. Skin and skull complete.
Type Series. 8 paratypes; B3854—B3856, B3858—
B3862; all represented by skin and skull, 3 females
and 5 males.
Synaptomys (Mictomys) innuitus
medioximus Bangs, 1900c
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) au Loup. 15 April 1899.
Collector. E. Doane. Original number 7.
Condition. Skin complete. Skull slightly damaged
(hole in left parietal).
Type Series. 1 paratype; B3972, skin and skull,
adult male.
Comments. S. b. medioximus was retained as a val-
id subspecies by Hall (1981: 834).
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
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 of 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 mearnsi Heller, 1910. See
Hubert (1978: 51).
Holotype. MCZ 31475. Skin and skull. 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 STEATOMYS 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. Skin and skull complete.
Type Series. 14 paratypes; MCZ 44214-44216,
4421844228; all represented by skin and skull, 9
males and 5 females.
LI 7
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., 58: 331, July.
=Tatera valida kempi Wroughton, 1906.
See Bates (1988: 277).
Holotype. MCZ 14491. Skin and skull. Adult female.
Locality. Sudan: Blue Nile, north of (Er) Roseires,
Aradeiba. 22 January 1913.
Collector. G. M. Allen and J. C. Phillips. Original
number 69. .
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Tatera soror G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 58: 333, July.
= Tatera valida kempi Wroughton, 1906.
See Bates (1988: 277).
Holotype. MCZ 14492. Skin and skull. Adult female.
Locality. Sudan: Blue Nile, Fazogli. 16 January
1913.
Collector. G. M. 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.
Type Series. Holotype only.
Subfamily MURINAE Illiger, 1811
Genus APODEMUS Kaup, 1829
Apodemus mystacinus euxinus G. M.
Allen, 1915b
Bull. Mus. Comp. Zool., 59: 11,
February.
=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
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.
Type Series. Holotype only.
Genus DASYMYS Peters, 1875
Dasymys incomtus alleni Lawrence and
Loveridge, 1953
Bull. Mus. Comp. Zool., 110: 53, June.
Holotype. MCZ 26322. Skin and skull. Adult male.
Locality. Tanganyika Territory (=Tanzania):
(Mbeya), Iolo, 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 skin
and skull, in the MCZ.
Comments. Retained as a valid subspecies by An-
sell (1978: 83).
Genus GRAMMOMYS Thomas, 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 HYLOMYSCUS Thomas, 1926
Hylomyscus alleni simus G. M. Allen and
Coolidge, 1930
In Strong, The African Republic of
Liberia and the Belgian Congo, 2: 599,
October.
=Hylomyscus alleni (Waterhouse, 1838).
See Heim de Balsac and Aellen (1965:
TID"
Holotype. MCZ 24028. Skin and skull. Adult male.
Locality. Liberia: Merikay. 13 September 1926.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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 MELOMYS Thomas, 1922
Melomys levipes stevensi Rummler, 1935
Z. Saugetierkunde, 10: 109, 31
December.
=Melomys mollis Thomas, 1913. See
Flannery (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 skin and skull, 4
females and 7 males.
Melomys moncktoni alleni Rummler, 1935
Z. Séugetierkunde, 10: 112, 31
December.
=Melomys rubex alleni Rummler, 1935.
See Tate (1951: 299).
Holotype. MCZ 29902. Skin and skull. Adult female.
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. 1 paratype; MCZ 29900, skin and
skull, adult male.
Comments. Flannery (1995a: 308-309) did not list
subspecies of M. rubex, noting, “It is a very 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’yMuss Comp. ZooleiSO 199!
February.
=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, Idjwi Island, Upper —
Mulinga, 6,500 ft (1,983 m). 24 February 1939.
Collector. A. Loveridge.
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes; MCZ 40746,
skull (no longer in MCZ); MCZ 40747,
skull, female.
skin and
skin and
Leggada gerbillus G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. Zool.,
February.
=Mus tenellus (Thomas, 1903). See
Musser and Carleton (1993: 629).
Holotype. MCZ 26586. Skin and skull. Adult male.
Locality. pei Territory (=Tanzania): Do-
doma, Ugogo, 3,700 ft (1, 129 m). 23 December
1929.
Collector. A. Loveridge.
Condition. Skin complete. Skull partial (right jugal
missing). Mandible disarticulated and angular pro-
cess of right ramus chipped.
Type Series. Holotype only.
Tee Aes.
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-
mone Mandible disarticulated.
Type Series. There is a series of paratypes in the
AMNH, anal 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 MYOMYS Thomas, 1915
Praomys fumatus oweni Setzer, 1956
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.
119
Collector. J. S. Owen. Original number 1030.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Genus NIVIVENTER Marshall, 1976
Epimys zappeyi G. M. Allen, 1912b
Mem. Mus. Comp. Zool., 40: 225,
August.
=WNiviventer confucianus (Milne-Edwards,
1871). See Musser and Carleton (1993:
683):
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 1908S.
Collector. W. R. Zappey. Original number 305.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Genus OENOMYS Thomas, 1904
Oenomys hypoxanthus talangae Setzer,
1956
Proc. U.S. Nat. Mus.,
November.
106: 505, 28
Holotype. MCZ 45315. Skin and skull. Adult female.
Locality. Anglo- Egyptian Sudan (=Sudan): Equa-
toria Province, Imatong Mountains, Talanga For-
est, 3,000 ft (915 m). 10 July 1950.
Collector. J. S. Owen. Original number 1338.
Condition. Skin and skull complete.
Type Series. | paratype; MCZ 45284,
skull, male.
Genus PRAOMYS Thomas, 1915
Praomys tullbergi melanotus G. M. Allen
and Loveridge, 1933
Bull. Mus. Comp. Zool.,
February.
=Praomys delectorum (Thomas, 1910).
See Musser and Carleton (1993: 642).
Holotype. MCZ 26287. Skin and skull. Adult male.
Locality. Tanganyika Territory (=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,
26256, 26288—26293, 26295-26297, 26387-26394,
26411, 26497, 26498: all represented by skin and
skull.
skin and
Tex Alley
120
Comments. Van der Straeten and Dieterlen (1987:
9) and Van der Straeten and Dudu (1990: 81) treat-
ed melanotus as a species within the Praomys de-
lectorum species complex.
Genus PSEUDOHYDROMYS Rummler,
1934
Pseudohydromys murinus Rummler, 1934
Z. Siiugetierkunde, 9: 48, 30 December.
Holotype. MCZ 29904. Skin 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. Skin complete, with small bald spots.
Skull complete. Mandible disarticulated.
Type Series. Holotype only.
Comments. Type species of Pseudohydromys Rii-
mmier, 1934, Considered a valid species by Flan-
nery (1995a: 255), Musser and Carleton (1993:
644), and Nowak (1999: 1612).
Genus STENOMYS Thomas, 1910
Stenomys niobe stevensi Rummler, 1935
Z. Séugetierkunde, 10: 117, 31
December.
=Stenomys niobe niobe (Thomas, 1906).
See Flannery (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 January 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. Siugetierkunde, 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-
nery (1995a: 346).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus THAMNOMYS Thomas, 1907
Thamnomys venustus kivuensis G. M.
Allen and Loveridge, 1942
Bull. Mus. Comp. Zool., 89: 192,
February.
Holotype. MCZ 39151. Skin and skull. 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.
Type Series. Holotype only.
Subfamily NESOMYINAE Forsyth Major,
1897
Genus NESOMYS Peters, 1870
Nesomys lambertoni G. Grandidier, 1930c
Bull. Acad. Malgache, n. ser., 11: 95 (for
1928).
Holotype. MCZ 45941. Skull and postcranial skele-
ton. Adult male.
Locality. Malagasy Republic (=Madagascar): west
coast, vicinity of Maintirano.
Collector, No collection data available. Received by
G. Grandidier from Académie Malgache, 1928.
Condition. Skull partial (left zygomatic arch miss-
ing, right jugal missing, several small holes). Post-
cranial skeleton complete (small foot bones pre-
sumably preserved in skin).
Type Series. 2 paratypes, 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,
February.
=Otomys lacustris G. M. Allen and
Loveridge, 1933. See Dieterlen and Van
der Straeten (1992: 385).
Holotype. MCZ 26358. Skin and skull. Adult female. |
Locality. Tanganyika Territory (=Tanzania): north —
end of Lake Nyasa, Ukinga Mountains, Madehani, —
7,000 ft (2,135 m). 21 February 1930.
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
Collector. A. Loveridge.
Condition. Skin complete. Skull 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.
Holotype. 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. Skin 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 by Ansell (1978: 78).
Subfamily SIGMODONTINAE Wagner,
1843
Genus ISTHMOMYS Hooper and Musser,
1964
Megadontomys flavidus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 27, April.
=Isthmomys 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.
121
Collector. W. W. Brown, Jr. Original number 28.
Condition. Skin complete, with small bald spots.
Skull complete. Mandible disarticulated.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. Type species of Isthmomys 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 January.
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, Jr. 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. |. abbreviata was retained as a valid
subspecies by Hall (1981: 755).
Neotoma bella Bangs, 1899j
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 angular process chipped.
IY Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Type Series. Holotype only.
Neotoma distincta Bangs, 1903a
Proc. Biol. Soc. Washington, 16: 89, 25
ijjunmies
=Neotoma mexicana distincta Bangs,
1903. See Hall (1955: 329).
Holotype. B9819. Skin and skull. Adult male.
Locality. Mexico: Vera Cruz, near Jalapa, Texolo. §
March 1899.
Collector. S. N. Rhoads. Original number 295.
Condition. Skin and skull complete.
Type Series. 4 paratypes; B9818, skin 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, 1898b
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, skin and
skull, adult male, and B2873, skin and skull, adult
female.
Comments. Retained as a valid subspecies by Hall
(1981: 749).
Genus NYCTOMYS de Saussure, 1860
Nyctomys nitellinus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 30, April.
=Nyctomys sumichrasti 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 February 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, skin 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,
February.
=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 Series. 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. BS101. Skin and skull. Adult female.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 24
March 1898.
Collector. W. W. Brown, Jr. Original number 89.
Condition. Skin complete. Skull partial (right jugal
missing).
Type 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, 1900
Oryzomys navus Bangs, 1899a
Proc. Biol. Soc. Washington, 13: 9, 31
January.
=Oligoryzomys fulvescens (Saussure,
1860). See Carleton and Musser (1989:
70).
Holotype. BS107. 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. Brown, Jr. Original number 154.
Condition. Skin and skull complete. Mandible dis- _
articulated.
Type Series. 9 paratypes; BS223-B8231; all repre-
sented by skin and skull, except B8228 (skin only);
2 females and 7 males. B8224 is now in the USNM, °
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
and B8223, B8226, B8230, and B8231 are now in
the AMNH.
Comments. Oryzomys navus is the type species of
Oligoryzomys 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 ORYZOMYS 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 paratypes; MCZ 10325, skin and
skull, adult female; MCZ 10326, skin and skull,
adult female; MCZ 10340, skin 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.
Locality. (United States): Florida, Monroe County,
Cape Sable. 17 April 1895.
Collector. C. L. Brownell. Original number 148.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
123
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, skin,
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 Gloger, 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 individuals.” The measurements 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
men discussed and described by measurement in
the original publication and the only adult speci-
men mentioned, is here designated as lectotype. P.
gossypinus was considered a valid species by Mus-
ser and Carleton (1993: 730) and Nowak (1999:
1361).
Hesperomys leucopus arcticus Mearns,
1890
Bull. Amer.
February.
Name preoccupied by Hesperomys
arcticus Coues, 1877
Mus. Nat. Hist., 2: 285, 21
Peromyscus maniculatus borealis Mearns,
1911
Proc. Biol. Soc. Washington, 24: 102, 15
May. (Replacement name for
Hesperomys 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.
Condition. Skin complete. Skull partial (left jugal
and process of squamosal missing, right process of
squamosal missing, basioccipital and palatine miss-
ing). Mandible disarueulated and both angular pro-
cess missing.
Type Series. Holotype 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).
Syntype. MCZ 5528. Skin and skull.
Locality. Nebraska (now Wyoming):
County), Deer Creek. 19 January 1860.
Collector. F. V. Hayden. Original no. 80. Formerly
USNM 4310.
Condition. Skin 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. | other syntype, present location un-
known, formerly in the USNM.
Comments. Hesperomys sonoriensis nebrascensis
(Converse
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Baird, 1858 is a nomen nudum. Osgood (1909: 78)
and Mearns (1911: 102) designated Deer Creek,
Nebraska, as the type locality. Jones (1958: 107—
111) reviewed the systematic history of nebrascen-
sis, and Jones and Mursaloglu (1961: 101-103) re-
ported the discovery of one oe the two original syn-
types in the MCZ. The location of the other syn-
type, 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. Skin 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 J. A. Allen, 1895.
See Schmidly (1973: 125).
Holotype. 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 skull complete. Mandible dis-
articulated.
Type Series.
adult female.
1 paratype; B5484, skin and skull,
Peromyscus cacabatus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 29, April.
= Peromyscus mexicanus (Saussure,
1860). See Huckaby (1980: 15).
Holotype. MCZ 10225. Skin 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
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 skull. Adult female.
Locality. (Canada): Nova Scotia, James River. 8
August 1894.
Collector. C. H. Goldthwaite.
Condition. Skin 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 P. 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.
125
Peromyscus gossypinus nigriculus Bangs,
1896e
Proc. Biol. Soc. Washington, 10: 124, 5
November.
=Peromyscus gossypinus gossypinus
(LeConte, 1853). See Osgood (1909: 136).
Holotype. B2731. Skin and skull. Adult female.
Locality. (United States): Louisiana, Plaquemines
Parish, Burbridge. 30 January 1895.
Collector. F. L. Small. Original number 1547/156.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a very 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
February 1895.
Collector. O. Bangs.
Condition. Skin and skull complete.
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
February.
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 valid subspecies by Hall
(1981: 685).
Peromyscus leucopus fusus Bangs, 1905a
Proc. New England Zool. Club, 4: 13, 28
February.
Holotype. B9737. Skin and skull. Adult male.
Locality. (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
(1981: 686).
Peromyscus oreas Bangs, 1898f
Proc. Biol. Soc. Washington, 12: 84, 24
March.
Holotype. B3696. Skin and skull. Adult female.
Locality. Canada: British Columbia, near boundary
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. Skin 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 February
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, 1896.
Peromyscus subgriseus baliolus Bangs,
1898n
Science, my ser.,.6: Zila. lOyAueust
(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: 2015 V5
March.
=Peromyscus polionotus rhoadsi 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. Skin and skull 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 January.
=Peromyscus maniculatus saturatus
Bangs, 1897. See Osgood (1909: 61).
Holotype. B2581. Skin and skull. Adult male.
Locality. Canada: British Columbia, Saturna Is-
land. 31 January 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 *¢ Helgen and McFadden 1
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).
Holotype. 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. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. R. s. vulcanius was retained as a valid
subspecies by Hall (1981: 644).
Reithrodontomys 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 February 1898.
Collector. T. Surber. Original number 466.
Condition. Skin 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 SCOTINOMYS Thomas, 1913
Akodon teguina apricus Bangs, 1902b
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 (=Bajo Bo-
quete), 4,000 ft (1,220 m). 24 February 1901.
ho
~
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, skin and skull, adult female.
Comments. S. t. apricus was retained as a valid sub-
species by Hooper (1972: 21) and Hall (1981: 734).
Akodon xerampelinus Bangs, 1902b
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. Skin 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, 1902b
Bull. Mus. Comp. Zool., 39: 32, April.
= Sigmodon hispidus borucae J. A. Allen,
1897. See Hall and Kelson (1959: 672).
Holotype. MCZ 10288. Skin and skull. Adult male.
Locality. Panama: Chiriqui, 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.”
Type 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
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 18101, skin and
skull, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 737).
Sigmodon hispidus furvus Bangs, 1903b
Bull. Mus. Comp. Zool., 39: 158, July.
Holotype. MCZ 10665. Skin and skull. Male.
Locality. Honduras: Atlantida, La Ceiba. 16 Janu-
ary 1902.
Collector. W. W. Brown, Jr. Original number 4.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 738). Sigmodon hispidus fervidus Lydekker,
1904 (p. 34) was a misspelling and thus accidental
renaming of Sigmodon hispidus furvus 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. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B8250, skin and _ skull,
adult male.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus THOMASOMYS 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. Skin and skull. Adult male.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Paramo de Macotama, 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). Mandible disarticulated.
Type Series. 4 paratypes; B8248, juvenile male,
B8345, adult female, B8346, adult female, B8347,
adult male; all represented by skin 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 J. 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. Skin 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
Condition. Skin 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.
Se eg Cet 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. Skin and skull complete.
Type Series. 2 paratypes; MCZ 17607, skin and
skull, adult female; and a specimen in the USNM,
number 125434.
Comments. Rosevear considered G. h.
“possibly valid as a race” (1969: 501).
argen teus
Claviglis soleatus collaris G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. Zool., 75:
February.
= Graphiurus lorraineus Dollman, 1910.
See Holden (1993: 764).
122,
Mas MCZ 26373. Skin and skull. Adult female.
Locality. Tanganyika Territory (=Tanzania): (Irin-
ga), north end Lake Nyasa, Ukinga Mountains, Ma-
dehani, 7,000 ft (2,135 m). 24 February 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.
=Graphiurus 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
a aso Nyiro River (=Ewaso Ngiro), 4,000 ft (1,220
m). 25 July 1909.
Gollector. 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 skull, adult female.
129
Graphiurus schwabi G. M. Allen, 1912a
Bull. Mus. Comp. Zool., 54: 441, April.
= Graphiurus 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 CRYPTOMYS Gray, 1864
Cryptomys hottentotus occlusus G. M.
Allen and Loveridge, 1933
Bull. Mus. Comp. Zool., 75: 125,
February.
=Cryptomys hottentotus whytei (Thomas,
1897). See Honeycutt et al. (1991: 51).
Holotype. MCZ 26557. Skin and skull. Adult male.
Locality. Tanganyika Territory (=Tanzania): (Irin-
a Ugungwe Mountains, Kigogo, 6,000 ft (1,830
m) 13) Janu: ary 1930.
@pllcciar A. Lov eridge.
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
Heterocephalus stygius G. M. Allen,
1912a
Bull. Mus. Comp. Zool., 54: 444, April.
= Heterocephalus glaber Ruppell, 1842.
See Hollister (1919: 160).
Holotype. MCZ 12470. Body in alcohol, skull extract-
ed. Adult female.
Locality. British East Africa (
Guaso Nyiro (
August 1909.
Collector. G. M. Allen.
Condition. Alcoholic.
disarticulated.
Type Series. Holotype only.
=Kenya): northern
=Ewaso Ngiro), Neumann’s Boma. 6
Skull complete. Mandible
130
Family ERITHIZONTIDAE Bonaparte,
1845
Genus ERITHIZON F. Cuvier, 1822
Erethizon dorsatus picinus Bangs, 1900c
Proc. New England Zool. Club, 2: 37, 20
September.
=Erithizon dorsatum dorsatum (Linnaeus,
1758). See Harper (1961: 90).
Holotype. B8839. Skin and skull. Adult male.
Locality. (Canada): Labrador, Strait of Belle Isle,
Lance (=Lanse) au Loup. 16 F ebruary 1899.
Collector. E. Doane.
Condition. Skin and skull complete.
Type Series. 15 paratypes; B8832—B8838, B8840—
B8847; all represented by skin and skull, 8 females
and 7 males. Bangs had also a skull of picinus
(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, 1901b
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, 1898k
Proc. Biol. Soc. Washington, 12: 163, 10
August.
=Dasyprocta punctata colombiana (Gray,
1898). See Cabrera (1961: 589).
BS8008. Skin and skull. Adult female.
Magdalena, Santa Marta. 6
Holotype.
Locality. Colombia:
January 1898.
Collector. W. W. Brown, Jr. Original number 37.
Condition. Skin and skull complete.
Type Series. | paratype; B8113, skin and skull, ju-
venile male.
Comments. D. p. colombiana was retained as a val-
id subspecies by Cabrera (1961: 589).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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. Skin and skull (and atlas).
Adult female.
Locality. Guadeloupe Island: Goyave. 22 August
1914.
Collector. G. K. Noble.
Condition. Skin complete. Skull partial (bone miss-
ing from nasal and frontal).
Type Series. 1 paratype; MCZ 15937,
skull, subadult female.
skin and
Dasyprocta punctata nuchalis Goldman,
A Oi7,
Proc. Biol. Soc. Washington, 30: 113, 23
May.
Holotype. MCZ 10081. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Divala. 30 November
1900.
Collector. W. W. Brown, Jr. Original number 17.
Condition. Skin and skull comple te.
Type Series. 4 paratypes; MCZ 10080, skin and
skull, adult male: 10084, skull: 10175, skin and
skull, adult female; 10176, skin and skull, adult
male.
Comments. Retained as a valid subspecies by Hall
(1981: S60).
Family AGOUTIDAE Gray, 1821
Genus AGOUTI Lacépede, 1799
Agouti paca virgatus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 47, April.
Holotype. MCZ 10079. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Divala. 16 December
1900.
Collector. W. W. Brown, Jr. Original number 21.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 858) and Pérez (1992: 1).
Family OCTODONTIDAE Waterhouse,
1839
Genus TYMPANOCTOMYS Yepes, 1941
Octomys barrerae Lawrence, 1941
Proc. New England Zool. Club, 18: 43,
28 January.
= Tympanoctomys barrerae (Lawrence,
1941). See Yepes (1942: 75).
Holotype. MCZ 39716. Skin and skull. Adult male.
Locality. Argentina: Mendoza Province, La Paz.
April 1939.
TYPE SPECIMENS OF RECENT MAMMALS ° Helgen and McFadden
Collector. J. M. de la Barrera. Original number
A381.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Type species of Tympanoctomys Yepes,
1941. T. barrerae was considerd a valid species by
Woods (1993: 789) and Nowak (1999: 1683).
Family ECHIMYIDAE Gray, 1825
Genus BOROMYS Miller, 1916
Boromys torrei G. M. Allen, 1917a
Bull. Mus. Comp. Zool., 61: 6, January.
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 zygomatic arch, pm‘ and
alveolar row of right side, m! 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 the 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 number 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 J. 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. Brown, Jr. Original number 130.
Condition. Skin and skull complete. Right dentary
broken at pm.
e331
Type Series. There is a large series of paratypes in
the MCZ.
Comments. P. s. burrus was retained as a valid sub-
species by Hall (1981: 873).
Proechimys gorgonae Bangs, 1905b
Bull. Mus. Comp. Zool., 46: 89, June.
Holotype. MCZ 10828. Skin 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.
Type 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
Univa Kansas Bulbl) Mus! Nataelelist ml:
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 (=Tauary), 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 skin and skull.
Comments. Retained as a valid subspecies by Ca-
brera (1961: 521). The use of the specific name
cayennensis over guyannensis follows Woods
(i993 7795):
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. Skin complete. Skull partial (bone miss-
ing from palatine to occipital).
Type Series. 1 paratype; MCZ 10997, skin and
skull.
Comments. Retained as a valid subspecies by Hall
(1981: 863).
132
Genus GEOCAPROMYS Chapman, 1901
Geocapromys cubanus G. M. Allen, 1917a
Bull. Mus. Comp. Zool. 61: 9, January.
= 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 of cubanus is stored
in the Vertebrate Paleontology Department of the
MCZ.
Geocapromys ingrahami abaconis
Lawrence, 1934
Occas. Pap. Boston Soc. 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 paratype 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.
Type 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) under the genus
Capromys. Woods (1993: 800) employed the genus
Geocapromys. G. i. irrectus is extinct. The type ma-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
terial of irrectus is stored in the Vertebrate Pale-
ontology Department of the MCZ.
Genus MESOCAPROMYS Varona, 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.
Type 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
Barus, 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, 1964
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 slightly along the posterolabial
wall.”
Type Series. Holotype only.
Comments. Considered a valid species by Woods
(1993: 804) and Nowak (1999: 1708). Complete
cranial and dentary material of araewm has been
collected in Haiti and deposited in the Florida Mu-
seum of Natural History (Woods 1993: 804). This
species is extinct. This specimen is stored in the
Vertebrate Paleontology Department of the MCZ.
o/
TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden
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 1030.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 7390-7392; all rep-
resented by skin 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, 1899g
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 skin
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 LEPUS 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.
133
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 Henry (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. Skin 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. Skin 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. Skin and skull complete.
Type Series. 2 other adult syntypes; USNM 4262/
38001, skin and skull, male: USNM 4263/4273,
skin and skull; 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
Lepus bairdi cascadensis Nelson, 1907
Proc. Biol. Soc. Washington, 20: 87, 11
December.
=Lepus americanus cascadensis Nelson,
1907. See Racey and Cowan (1936:
H1S8).
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. Skin 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 SYLVILAGUS 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. Brown, 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 palustris paludicola (Miller
and Bangs, 1894). See Nelson (1909:
269).
Holotype. B1451. Skin and skull. Adult female.
Locality. (United States): Florida, (Citrus County),
Fort Island, near Crystal River. 25 January 1894.
Collector. F. L. Small. Original number 1150.
Condition. Skin and skull complete.
Type Series. 3 paratypes; B1452, skin and skull,
adult male; B1453, skin and skull, adult male;
B1454, skin and skull, adult female.
Comments. S. p. paludicola was retained as a valid
subspecies by Hall (1981: 299) and Chapman and
Willner (1981: 1).
Lepus sylvaticus alacer Bangs, 1896h
Proc. Biol. Soc. Washington, 10: 136, 28
December.
= Sylvilagus floridanus alacer (Bangs,
1896). See Lyon (1904: 336).
Holotype. B5480. Skin and skull. Adult female.
Locality. (United States): Indian Territory
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
(=Oklahoma), (Adair County), Stilwell. 14 August
1896.
Collector. T. Surber. Original number 65.
Condition. Skin 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).
Holotype. B2407. Skin and skull. Adult female.
Locality. (United States): Connecticut, New Lon-
don County, Liberty Hill. 6 November 1894.
Collector. O. Bangs.
Condition. Skin 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:
LW)
Order SCANDENTIA Wagner, 1855
Family TUPAIIDAE Gray, 1825
Genus TUPAIA Raffles, 1822
Tana tana griswoldi Coolidge, 1938
Proc. New England Zool. Club, 17: 45, 6
May.
=Tupaia tana paitana (Lyon, 1913). See
Medway (1965: 76).
Holotype. MCZ 36416. Skin, skull, 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. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
TYPE SPECIMENS OF RECENT MAMMALS *¢ Helgen and McFadden
Order PRIMATES Linnaeus, 1758
Family INDRIDAE Burnett, 1828
Genus PROPITHECUS Bennett, 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-
ary), [fotaka, vicinity of Ambovombe. Probably
1928.
Collector. M. Perrier. Original number 1. Grandi-
dier Collection.
Condition. Skin and skull complete. Postcranial
skeleton partial (right humerus broken, foot bones
left in skin).
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. Geoffroy, 1795
Cheiromys madagascariensis laniger G.
Grandidier, 1930d
Bull. Acad. Malgache, n. ser., 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
Académie 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 Académie 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).
Si)
Family GALAGONIDAE Gray, 1825
Genus GALAGOIDES A. Smith, 1833
Galago demidovii orinus Lawrence and
Washburn, 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 Territory (=Tanzania): Mor-
ogoro, Uluguru Mountains, Bagilo, 5,000 ft (1,525
m). 17 September 1926.
Collector. A. Loveridge.
Condition. Skin and skull complete.
Type Series. Holotype only.
Family CEBIDAE Bonaparte, 1831
Genus ALOUATTA Lacépéde, 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 skull. 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.
Type Series. Holotype 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. Barbour. Original number 4.
Condition. Skin and skull complete.
Type Series. Lawrence mentions examining 19
specimens of trabeata and lists them by locality
(1933: 330). 9 of these are in the MCZ; other than
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
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 lIlliger, 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 155.
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,
Harvard, 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 |. Geoffroy, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Locality. Gabon: Gabon Estuary, Mpongwe coun-
try. 1847.
Collector. T. S. Savage. Original number 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 pelvis, 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
Br the original four to be noted in Wyman’s per-
sonal notebook of osteology (unpublished, now in
the library of the Boston Museum of Science) and
the only original specimens of which there is any
record at Fil 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 revision 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
(1993: 276) and Nowak (1999: 618).
Order LIPOTYPHLA Haeckel, 1866
Family NESOPHONTIDAE Anthony, 1916
Genus NESOPHONTES Anthony, 1916
Nesophontes micrus G. M. Allen, 1917a
Bull. Mus. Comp. Zool., 61: 5, January.
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 1
Condition. Condition as originally described—
“posterior half of the right ramus, containing a part
of pm, m,, ms, and the roots of m;.”
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 SOLENODON Brandt, 1833
Antillogale marcanoi Patterson, 1962
Breviora, Mus. Comp. Zool., 165: 3, 22
August.
= Solenodon marcanoi (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 P;-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. Sh @
oN)
~l
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, Martha’s Vineyard, West Tisbury. 25 June
1899.
Collector. O. Bangs. Original number 2.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 9 paratypes, B9725, B9726, B9728—
B9734, all represented by skin and skull, 3 females
and 6 females.
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. B97O5. 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, B97O06,
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 CROCIDURA Wagner, 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. Skin and skull complete.
Type Series. Setzer mentions that he examined 18
specimens of tephragaster, 8 of which are in the
MCZ.
138
Crocidura hildegardeae phaios Setzer,
1956
ErOcw Wr SeaNat,
November.
Mus., 106: 460, 28
Holotype. MCZ 45855. 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 hilde gardeae by
Hutterer (1993: 87).
Genus CRYPTOTIS Pomel, 1848
Cryptotis avia G. M. Allen, 1923a
Proc. New England Zool. Club, 8: 37, 12
February.
=Cryptotis thomasi (Merriam, 1897). See
Woodman (1996: 414).
Holotype. MCZ 20091. Skin and skull. Adult.
Locality. Colombia: (probably Cundinamarca), E]
Verjon (see coments). October 1922.
Collector. N. Maria.
Condition. Skin 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 av ailable 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., 3
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 Territory (=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.
8: 417, 23
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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 sere gloveralleni Jackson, 1926
J. Mammal., 7: 57, 15 February.
(once ee 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;
USNM in 1922.
Comments. Retained as a valid subspecies by Hall
(1981: 41) and Beneski and Stinson (1987: 1).
B2053, exchanged to the
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. Skin 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,
April.
= Sorex caecutiens koreni G. M. Allen,
1914. See Yudin (1989: 281).
Holotype. MCZ 15085. Skin and skull. 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.
De OOo
TYPE SPECIMENS OF RECENT MAMMALS ° Helgen and McFadden
Type Series. 5 paratypes; MCZ 15003-15007, all
represented by skin 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, §
December.
Name preoccupied by Sorex macrourus
Lehmann, 1822.
Sorex dispar Batchelder, 1911
Proc. Biol. Soc. Washington, 24: 97, 15
May. (Replacement name for Sorex
macrurus Batchelder, 1896)
Holotype. MCZ 41744. Skin and skull. 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 locality 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
February.
= 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. Skin 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 daphaenodon sanguinidens G. M.
Allen, 1914. See Yudin (1989: 198).
Holotype. MCZ 15012. Skin and skull. Adult female.
Locality. (Russian Federation): northeastern Sibe-
139
ria, Nijni Kolymsk (=Nizhnekolymsk), near mouth
Kolyma River. 11 December 1911.
Collector. J. Koren. Original number 221.
Condition. Tail separate from skin. 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 SUNCUS Ehrenberg, 1832
Suncus ater Medway, 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,
February.
Holotype. MCZ 26754. Skin and skull. 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. Holotype only.
Comments. Retained as a valid subspecies by Heim
de Balsac and Meester (1977: 6).
140
Family TALPIDAE Fischer de Waldheim,
Ws y
Genus NEUROTRICHUS Gunther, 1880
Neurotrichus gibbsi hyacinthinus Bangs,
1897d
Amer. Nat., 31: 240, 1 March.
Holotype. B1240. Skin 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, 1804
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 February
1897.
Collector. O. Bangs. Original number 10.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 4 paratypes; B7193—B7196; all repre-
sented by skin 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, 1896h
Proc. Biol. Soc. Washington, 10: 138, 28
December.
= Scalopus aquaticus aereus (Bangs,
1896). See Miller (1912: 8).
Holotype. B5475. Skin and skull. Adult female.
Locality. (United States): Indian Territory
(=OkKahoma) (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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus SCAPANUS Pomel, 1848
Scapanus Californicus minusculus Bangs,
1899}
Proc. New England Zool. Club, 1: 70, 31
July.
= 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. l. 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-
bah), Mount Kinabalu, Luma Luma, 5,500 ft (1,678
m). 12 July 1937.
Collector. J. A. Griswold, Jr., Asiatic Primate Ex-
pedition. Original number 510.
Condition. Skin, skull, and posteranial skeleton
complete.
Type Series. 3 paratypes; MCZ 36583, skin, 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
In Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 38, November.
Holotype. MCZ 35159. Skin and skull. Adult male.
Locality. Philippines: Luzon, (Nueva Ecija), Rizal,
Montalban caves near Manila. 27 February 1937.
Collector. B. Lawrence. Original number 253.
Condition. Skin and skull complete.
Type Series. In the original description, Lawrence
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
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), Mount
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, 1777
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 Flannery (1995b).
Pteropus ariel G. 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 Expedition.
Condition. Right wing of skin 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).
141
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 February 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. Adult male.
Locality. Solomon Islands: Treasury (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. ser., 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
Académie Malgache, 1917.
Condition. 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 R. madagascariensis,
Bergmans (1977: 67) commented in error that the
holotype was in the Académie Malgache, Antana-
narivo, Madagascar.
142
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).
Holotype. 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. Washington, 27: 109, 10
July.
=Rhynchonycteris naso (Wied-Neuwied,
1820). See Sanborn (1937: 326).
Holotype. MCZ 13208. Skin and skull. Adult.
Locality. Mexico: 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
skull, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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: 63).
Nycteris nana tristis G. M. Allen and
Lawrence, 1936
Bull. Mus. Comp. Zool., 79: 47, January.
=WNycteris 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 February 1934.
Collector. A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Family RHINOLOPHIDAE Gray, 1825
Genus HIPPOSIDEROS Gray, 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.
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).
extracted but not
Skull
Hipposideros erigens Lawrence, 1939
In Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 56, November.
= Hipposideros 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, skin 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 * Helgen and McFadden
Hipposideros turpis Bangs, 1901a
Amer. Nat., 35: 561, 31 July.
= Hipposideros turpis turpis Bangs, 1901.
See Hill (1963: 94).
Holotype. MCZ 10003. Skin and skull. Adult female.
Locality. (Japan): Ryukyu Islands, southern group
of Liu Kiu Islands, Ishigaki Island. 10 May 1899.
Collector. 1. Zensaku.
Condition. Skin complete. Skull partial (occipital
region missing).
Type Series. 2
skull, adult female;
adult male.
Comments. H. turpis was considered a valid species
by Koopman (1993: 175) and Nowak (1999: 334).
Genus RHINOLOPHUS Lacépéde, 1799
Rhinolophus megaphyllus ignifer G. M.
Allen, 1933
J. Mammal., 14: 149, 15 May.
= Rhinolophus megaphyllus megaphyllus
Gray, 1834. See Koopman (1984: 9).
Holotype. MCZ 29078. Skin and skull. Adult male.
Locality. Australia: Queensland, Cape York, Coen.
12 June 1932.
Collector. P. J. Darlington, Jr.,
Expedition. Original number 185.
Condition. Slani: and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 29079,
skull, adult male.
paratypes; MCZ 10002, skin and
MCZ 10004, skin and _ skull,
Harvard Australian
skin and
Rhinolophus philippinensis alleni
Lawrence, 1939
In Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 46, November.
Holotype. MCZ 35097. Skin 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 paratypes;
skull, adult female; MCZ
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
January.
Holotype. MCZ 45080. Muminy.
Locality. Malagasy Republic (=Madagascar): (An-
tsiranana), Diego-Suarez (=Antsiranana). 1910.
MCZ
35099,
35098, skin and
skin and _ skull,
143
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
Chilonycteris 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).
Chilonycteris torrei G. M. Allen, 1916a
Proc. New England Zool. Club, 6: 4, 8
February.
=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.,
May.
=Ametrida centurio Gray, 1847. See
Peterson (1965: 5).
Holotype. MCZ 11274. Body in alcohol, skull extract-
ed. Adult male.
26: 240, 16
144
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.
Type Series. Holotype only.
Genus EROPHYLLA Miller, 1906
Erophylla sezekorni syops G. M. Allen,
1917c
Proc. Biol. Soc. Washington, 30: 167, 23
October.
Holotype. MCZ 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 exchange to the USNM.
Comments. Retained as a valid subspecies by Hall
(1981: 171) and Koopman (1994: 79).
Genus GLOSSOPHAGA E. Geoffroy,
1818
Glossophaga longirostris Miller, 1898
Proc. Acad. Nat. Sci. Philadelphia, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
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 de Saussure,
1860
Vampyrops umbratus Lyon, 1902
Proc. Biol. Soc. Washington, 15: 151, 20
June.
=Platyrrhinus umbratus (Lyon, 1902). See
Koopman (1993: 191).
Holotype. B8180. Skin and skull. Adult male.
Locality. Colombia: (LaGuajira), San Miguel. 8
June 1898.
Collector. W. W. Brown, Jr. Original number 234.
Condition. Skin and skull complete.
Type Series. 2 paratypes; B8300, skin, male; B8301,
skin, 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
Comments. In the original description, H. Allen re-
ported 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 with the accession
catalogue of the MCZ (1931: 236-237). In support
of Allen’s decision, Rouk and Carter (1972: 4) stat-
ed, after examining the holotype of zarhinus, that
it is “quite like specimens of [Platyrrhinus] 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, 1900
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,
February.
=Mops condylurus leucostigma (G. M.
Allen, 1918). See Koopman (1994: 141).
Holotype. MCZ 16344. Skin and skull. Adult 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.
145
Mops angolensis orientis G. M. Allen and
Loveridge, 1942
Bull. Mus. Comp. Zool., 89: 166,
February.
=Mops condylurus orientis G. M. Allen
and Loveridge, 1942. See Koopman
(1994: 141).
Holotype. MCZ 38829. Skin and skull. Adult male.
Locality. Tanganyika Territory (=Tanzania): Mtwa-
ra, Ruvuma River, Kitaya, 300 ft (92 m). 3 April
1939.
Collector. A. Loveridge.
Condition. Skin and skull complete.
Type 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-
tablishment, 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 Flannery (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 EPTESICUS 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 Australian
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
537) considered darlingtoni to be a valid species of
Vespadelus, 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).
Holotype. MCZ 14206. Skin and skull. Adult male.
Locality. Vietnam: Tonkin, Lao-Kai (=Lao Cai). 3
January 1912.
Collector. Kobayashi Collection. Original number
14.
Condition. Skin complete. Skull partial (parietals
broken).
Type Series. Holotype only.
Comments. H. h. rufulus was retained as a valid
subspecies by Koopman (1994: 133).
Genus IDIONYCTERIS Anthony, 1923
Corynorhinus phyllotis G. M. Allen, 1916b
Bull. Mus. 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 Potosf. 24 March 1878.
Collector. E. Palmer.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. I. phyllotis was considered a valid spe-
cies by Koopman (1993: 205) and Nowak (1999:
457).
Genus LAS/URUS Gray, 1838
Atalapha brachyotis J. 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. Lasiurus 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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus MYOTIS 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. Skin 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. Skin 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 paratype, 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 the
FMNH, USNM, AMNH, and BMNH.
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
Genus NYCTICEIUS Rafinesque, 1819
Nycticeius africanus G. M. Allen, 1911b
Bull. Mus. Comp. Zool., 54: 328,
December.
=Nycticeius schlieffeni albiventer Thomas
and Wroughton, 1908. See Hayman and
Hill (1971: 36).
Holotype. MCZ 8272. Skin and skull. Male.
Locality. British oe Africa (=Kenya): Meru Riv-
er, effluent of northern Guaso Nyiro (=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.
= Pipistrellus rendalli phasma (G. M. Allen,
191); 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 notius G. M. Allen, 1908
Bull. Mus. Comp. Zool., 52: 46, July.
=Pipistrellus 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. Layard, August
1864.
Condition. Alcoholic, skull partial (right and left
zygomatic arches missing; supraoccipital chipped).
Type Series. Holotype only,
Genus PLECOTUS E. Geoffroy, 1813
Plecotus sacrimontis G. M. Allen, 1908
Bull. Mus. Comp. Zool., 52: 50, July.
=Plecotus 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.
147
Collector. A. Owston.
Condition. Alcoholic.
Type Series. Holotype only.
Comments. P. a. sacrimontis was retained as a valid
subspecies by Koopman (1994: 110).
Genus SCOTOPHILUS Leach, 1821
Scotophilus altilis G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 58: 350, July.
= Scotophilus leucogaster (Cretzschmar,
1826). See Koopman (1993: 227).
Holotype. MCZ 14463. Skin and skull. Adult male.
Locality. Sudan: Blue Nile, north of (Er) Roseires,
Aradeiba. 22 a ary 1913.
Collector. G. Allen, Phillips Sudan Expedition.
Original Mehta 73.
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 Spix, 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. Duryea.
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 PECARI Reichenbach, 1835
Tayassu crusnigrum Bangs, 1902b
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 (=Bajo Bo-
quete), 4,000 ft (1,220 m). 13 April 1901.
148
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. crusnigrum was retained as a valid
subspecies by Hall (1981: 1080) under the genus
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Locality. China: Kiangsu (=Jiangsu), Kiangyin, 80
miles (129 km) northwest of Shanghai. 7 April
1922.
Collector. F. R. Wulsin.
Condition. Skin and postcranial 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 Delphinus melas Schlegel, 1841,
Dicotyles.
Tayassu torvus Bangs, 1898k
Proc. Biol. Soc. Washington, 12: 164, 10
August. cs
5 I) eh the holotype of this new name is the same as that
=Pecari tajacu torvus (Bangs, 1898). See of Schlegel’s name, RMNH 23079.
comments.
Family CERVIDAE Goldfuss, 1820
Genus ODOCOILEUS 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-
a articulated.
Beluga declivis Cope, 1865 Type Series. 4 paratypes; B2391, adult male,
Proc. Acad. Nat. Sci. Philadelphia, 17: B2392, adult male, B2393, adult female, B2395,
27S. juvenile male, all represented by skin and skull.
=Delphinapterus leucas (Pallas, 1776). Comments. O. v. osceola was retained as a valid
Gan ee een (1966: aa ) i baka by Hall (1981: 1096) and Smith (1991:
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.
Holotype. BS038. Skin 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 Lacépede,
1804
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. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B9112, B8622, B8623, all
males represented by skin and skull.
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
ens WS CnOCAIEN A Pals Ue Odocoileus americanus borealis Miller,
Neomeris asiaeorientalis Pilleri and Gihr, 1900
1972 Bull. New York State Mus. 8: 83, 21
Invest. Cetacea, 4: 126. November.
=Neophocaena phocaenoides
asiaorientalis (Pilleri and Gihr, 1972). See
van Bree (1973: 17).
Holotype. MCZ 19998 (but see comments). Skull and
postcranial skeleton. Adult male.
=Odocoilus virginianus borealis Miller,
1900. See Trouessart (1905: 704).
Holotype. B4999. Skin and skull. Adult male.
Locality. (United States): Maine, (Hancock Coun-
ty), Bucksport. 12 December 1895.
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
Collector. A. G. Dorr.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. O. v. borealis was retained as a valid
subspecies by Hall (1981: 1092) and Smith (1991:
ib)
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 skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes, MCZ 18497, skin and
skull, juvenile male; MCZ 18060, skin and skull,
juvenile male.
Comments. Retained as a valid subspecies by Hall
(1981: 1093) and Smith (1991: 1).
Genus RANGIFER 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. Bryant.
Condition. Single antler; complete.
Type Series. Holotype only.
Rangifer terraenovae Bangs, 1896f
Preliminary Description of the
Newfoundland Caribou, Boston, p. 1, 11
November.
= Rangifer tarandus caribou (Gmelin,
1788). See Banfield (1962: 70).
Holotype. B3778. Skull and head skin. Adult male.
Locality. Canada: Newfoundland, Codroy. 8 Sep-
tember 1895
Collector. F. Doane.
Condition. Skull and head skin 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
149
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 DAMALISCUS Sclater and
Thomas, 1894
Damaliscus phillipsi Harper, 1939
Proc. Biol. Soc. Washington, 50: 90, 5
June.
=Damaliscus pygargus phillipsi 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 Harper 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 pygargus
over dorcas, see Rookmaaker (1991: 190).
Order CARNIVORA Bowdich, 1821
Family CANIDAE Fischer de Waldheim,
1817
Genus CANIS Linnaeus, 1758
Canis lupus beothucus 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 postecranial 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. |. beothucus became extinct around
1911. Retained as a valid subspecies by Hall (1981:
930).
150
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). PB. robustus is the type species of Pa-
chycyon J. A. Allen, 1885. Pachycyon is a synonym
of Canis Linnaeus, 1758, which is commonly over-
looked, for example, in Wozencraft (1993) 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 savannah of Bogota, 9,000
ft (2,745 m).
Colleeror N. Maria. Original number 25.
Condition. Skin and skull complete.
Type Series. 5 paratypes; MCZ 19849, skin and
skull, juvenile; MCZ 20097, skin 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, 1898h
Proc. Biol. Soc. Washington, 12: 93, 30
April.
=Cerdocyon thous aquilus (Bangs, 1898).
See Langguth (1969: 178).
Holotype. BSO001. 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.
adult female.
Comments. Retained as a valid subspecies by Berta
@LOS2 auld)»
1 paratype; BS002, skin and skull,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus UROCYON 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. 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).
Urocyon cinereoargenteus ocythous
Bangs, 1899h
Proc. New England Zool. Club,
June.
1: 43, 5
Holotype. B4290. Skin and skull. Adult female.
Locality. (United States): Wisconsin, Grant Coun-
ty, Platteville. 25 January 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 VULPES Frisch, 1775
Vulpes deletrix Bangs, 1898d
Proc. Biol. Soc. Washington, 12: 36, 24
March.
= Vulpes vulpes rubricosa Bangs, 1898.
See Churcher (1960: 359).
Holotype. B6967. Skin and skull. Adult female.
Locality. (Canada): Newfoundland, Bay St.
George. 24 April 1897.
Chllezor E. Doane.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Vulpes pennsylvanica vafra Bangs, 1897f
Proc. Biol. Soc. Washington, 11: 53, 16
March.
Name preoccupied by Vulpes vafer
Leidy, 1869.
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
Vulpes pennsylvanica rubricosa Bangs,
1898a
Science, n. ser., 7: 271, 25 February.
(Replacement name for Vulpes
pennsylvanica 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
skull, and B2002, skull.
Comments. V. v. rubricosa was retained as a valid
subspecies by Hall (1981: 939). Vulpes fulvus rub-
ricatus 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 URSUS Linnaeus, 1758
Ursus (Euarctos) americanus sornborgeri
Bangs, 1898)
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
151
true holotype 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 lotor (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-
tury (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.
Locality. (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 lotor (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
Family MUSTELIDAE Fischer de
Waldheim, 1817
Genus LONTRA Gray, 1843
Lutra degener Bangs, 1898d
Proc. Biol. Soc. Washington, 12: 35, 24
March.
=Lontra canadensis canadensis
(Schreber, 1776). See van Zyll de Long
@lIS72aS1))
Holotype. B6965. Skin 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, 1898b
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 County,
Micco. 17 March 1897.
Collector. F. R. Hunter.
Condition. Skin and skull complete.
Type Series. 4 paratypes; B4995, skin and skull,
adult female; B4998, skin and skull, adult male;
BG6092, skin and skull, adult male; B6093, skin 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. Skin 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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Mustela brumalis Bangs, 1898)
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 richardsonii 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 Cockerell, 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 February 1898.
Collector. A. C. Tyson. Original number 58.
Condition. Skin 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 history, 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 *¢ Helgen and McFadden
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 Columbia, source of Ket-
tle River, 7,500 ft (2,288 m). 10 September 1898.
Collector. A. C. Brooks. Original number 1368.
Condition. 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, 1898b
Proc. Boston Soc. Nat. Hist., 28: 229, 15
March.
= Mustela vison lutensis (Bangs, 1898).
See Hollister (1913: 474).
Holotype. B7225. Skin and skull. Adult male.
Locality. (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 Lariviére
(1999: 1).
Putorius (Arctogale) muricus Bangs, 1899}
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.
Condition. 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,
1899)
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, Weaverville. 10 July 1892.
153
Collector. J. S. Cairns. Original number 2214.
Condition. Skin 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, 1899i
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
February.
= Mustela nivalis rixosa (Bangs, 1896).
See Reichstein (1958: 169).
Holotype. B642. Skin and skull. Adult female.
Locality. Canada: Saskatchewan, Osler. 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
Type Series. | paratype; B3556, skin and skull, ju-
venile male.
Comments. M. v. energumenos was retained as a
valid subspecies by Hall (1981: 1001) and Lariviére
(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 January 1895.
Collector. F. L. Small. Original number 1439/54.
Condition. Skin and skull complete.
Type Series. 10 paratypes; B2752-B2761, all rep-
resented by skin and skull, 9 males and 1 female.
Comments. M. v. vulgivaga was retained as a valid
subspecies by Hall (1981: 1004) and Lariviére
(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. | paratype, BS8632 (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. Geoffroy 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. ;
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Condition. Skin and skull complete.
Type 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, 1896i
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,
Cartville. 25 May 1895.
Collector. F. L. Small. Original number 1842.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype, 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 ¢ Helgen and McFadden
Genus SPILOGALE Gray, 1865
Spilogale ambarvalis Bangs, 1898b
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 Micco. 30 Jan-
uary 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 Lavauden, 1929
Compt. Rend. Acad. Sci. Paris, 189: 198,
QS) July DOR
=Eupleres goudotii major Lavauden,
1929. See Albignac (1973: 23).
Syntypes. MCZ 45691: Skin, skull, and poster: anial
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, with bald
spot on dorsum and tail slightly damaged. Skull and
skeleton complete. Mandible eneeculated 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 Ewpleres ma-
jor. Albignac (1973: 23) wrote that these type spec-
imens were “introuvable [nowhere to be found].”
Family HERPESTIDAE Bonaparte, 1845
Genus GALIDICTIS |. Geoffroy, 1839
Galidictis grandidiensis [sic] Wozencraft,
1986
J. Mammal. 67: 561, 8 August.
= Galidictis grandidieri Wozencraft, 1986.
See Wozencraft (1987: 198).
Holotype. MCZ 45983. Skin, skull, and postcranial
skeleton. Adult.
155
Locality. Madagascar (no further data available).
The locality of the paratype, stored in the AMNH,
s “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 paratype; AMNH 100478, skin 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 Severtzov, 1858
Felis capensis phillipsi G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 58: 337, July.
=Leptailurus serval phillipsi (G. M. Allen,
1914). See comments.
Holotype. MCZ 14908. Skin and _ skeleton. Adult
male.
Locality. Sudan: Blue Nile, El Garef. 10 January
1913.
Collector. J. C. Phillips.
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-
tailurus follows Wozencraft (1993: 292). L. s. phil-
lipsi was retained as a valid subspecies by Smithers
(1975: 7).
Genus LYNX Kerr, 1792
Lynx (Cervaria) fasciatus oculeus Bangs,
1899e
Proc. New England Zool. Club, 1: 23, 31
March.
= Lynx rufus californicus Mearns, 1897.
See Grinnell and Dixon (1924: 346).
Holotype. B8633. Skin and skull. Adult male.
Locality. (United States): California, Marin Coun-
ty, Nicasio. 11 December 1898.
Collector. C. A. Allen. Original number 981.
Condition. Skin and skull complete.
Type Series. 1 paratype; B4789, skin 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
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 Elliot (1901: 296).
Holotype. B1190. Skin 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 bangsi Merriam, 1901
Proc. Washington Acad. Sci., 3: 595, 11
December.
=Puma concolor bangsi (Merriam, 1901).
See comments.
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).
Felis 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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Felis coryi Bangs, 1899b
Proc. Biol. Soc. Washington, 13: 15, 31
January.
=Puma concolor coryi (Bangs, 1899). See
comments.
Holotype. B7742. Skin and skull. Adult male.
Locality. (United States): Florida, Brevard County,
“wilderness back of Sebastian”. 1 January 1898
Collector. F. R. Hunter.
Condition. Skin and skull complete.
Type Series. 5 paratypes; B5489, adult female,
B5650, adult female, B6992, adult male, B7743,
adult female, B7744, juvenile female; all repre-
sented by skin and skull.
Comments. Nelson and Goldman (1929: 347) used
the name 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 PHATAGINUS Rafinesque, 1821
Phataginus tricuspis mabirae G. M. Allen
and Loveridge, 1942
Bull. Mus. Comp. Zool., 89: 178,
February.
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 yz
Comments. Retained as a valid subspecies by
Meester (1972: 2).
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INDEX
abaconis
Geocapromys, 132
abbotti
Myotis, 146
abbreviata
Neotoma, 121
aberti
Sciurus, 99
abietorum
Peromyscus, 125
ablutus
Leggada, 118
acadicus
Microtus, 115
Neosorex, 138
Zapus, 112
aequalis
Aethalops, 140
aereus
Scalops, 140
Scalopus, 140
aestuans
Guerlinguetus, 106
Sciurus, 106
Aethalops
aequalis, 140
alecto, 140
Aethoglis
argenteus, 129
hueti, 129
Aethosciurus
byatti, 106
laetus, 106
africanus
Nycticeius, 147
Afrosoricida, 101
Agouti
paca, 130
virgatus, 130
Agoutidae, 130
akka
Funisciurus, 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
Pteromys, 105
alecto
Aethalops, 140
alfari
Microsciurus, 105
alleni
Dasymys, 118
Heteromys, 112
Hylomyscus, 118
Lepus, 133
Liomys, 112
Macrotolagus, 133
Melomys, 118
Rhinolophus, 143
aloga
Blarina, 137
Alouatta
luctuosa, 135
palliata, 135
pigra, 135
trabeata, 135
alpinus
Sciuropterus, 104
altilis
Scotophilus, 147
amazonicus
Nectomys, 121
ambarvalis
Spilogale, 155
americana
Martes, 152
americanus
Euarctos, 151
Lepus, 133, 134
Odocoileus, 148
Ursus, 151
Ametrida
centurio, 143
minor, 143
ammodytes
Peromyscus, 125
anastasae
Peromyscus, 124, 125
Scalops, 140
Scalopus, 140
anchietae
Otomys, 120
anchises
Hylopetes, 105
Pteromys, 105
anetianus
Pteropus, 141
angolensis
Mops, 145
Antechinus
mayeri, 100
misim, LOO
naso, 100
Antillogale
marcanoi, 137
aorensis
Pteropus, 141
Aplodontia, 103
californica, 103
columbiana, 103
rainieri, 103
rufa, 103
Aplodontidae, 103
Apodemus
euxinus, 117
mystacinus, 17,
apricus
Akodon, 127
Scotinomys, 127
aquaticus
Scalopus, 140
aquilis
Cerdocyon, 150
Craseomys, 113
Urocyon, 150
aquilonius
Fiber, 116
Ondatra, 116
araeum
Plagiodonta, 132
araneus
Sorex, 138
arcticus
Hesperomys, 124
Lepus, 133
Rangifer, 149
Arctogale
longicauda, 153
muricus, 153
oribasus, 153
Arctomys
avarus, 105
flaviventer, 105
ignavus, 105
arenarius
Peromyscus, 126
argentatus
Peromyscus, 125
argenteus
Aethoglis, 129
Graphiurus, 129
ariel
Pteropus, 141
armatus
Spermophilus, 107
Artibeus
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden ill
femurvillosum, 144
lituratus, 144
palmarum, 144
Artiodactyla, 145
Arvicola
breweri, 114
longipilis, 114
riparia, 114
rufidorsum, 114
terraenovae, 114
Arvicolinae, 112
asiaeorientalis
Neomeris, 148
Neophocaena, 148
Atalapha
brachyotis, 146
ater
Suncus, 139
atrata
Martes, 152
Mustela, 152
atrodorsalis
Thomomys, 111
attwateri
Peromyscus, 124
aurita
Geogale, 101
Triaenops, 143
auritus
Plecotus, 147
aurora
Eothenomys, 113
Microtus, 113
austerulus
Sigmodon, 127
austini
Pteropus, 141
australis
Cryptogale, 101
Echymipera, 101
Reithrodontomys, 127
austrinus
Geomys, 110
avara
Marmota, 105
avarus
Arctomys, 105
avia
Cryptotis, 138
Mephitis, 154
bactrianus
Mus, 121
badius
Ictidomys, 108
Spermophilus, 108
bairdii
Lepus, 133, 134
172 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Mus, 98
Peromyscus, 98
baliolus
Peromyscus, 126
baluensis,
Callosciurus, 103
bangsi
Felis, 156
Glaucomys, 104
Perognathus, 112
Puma, 156
Sciuropterus, 104
Vulpes, 151
bangsii
Lepus, 133
barbouri
Otomys, 121
barrerae
Octomys, 130
Tympanoctomys, 130
Bathyergidae, 129
bella
Neotoma, 121
bellus
Peromyscus, 124
Beluga
concreta, 145
declivis, 148
beothucus
Canis, 149
bicolor
Crocidura, 137
Hipposideros, 142
Oecomys, 122
Blarina
aloga, 137
brevicauda, 137
compacta, 137
borealis
Lasiurus, 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
breweri
Arvicola, 114
Microtus, 114
brochus
Syntheosciurus, 108
browni
Microsciurus, 105
Sciurus, 105
brumalis
Martes, 152
Mustela, 152
bufo
Leggada, 118
Mus, 118
burrus
Proechimys, 131
byatti
Aethosciurus, 106
Paraxerus, 106
caboti
Rangifer, 149
cacabatus
Peromyscus, 124
caecator
Castor, 109
caecutiens
Sorex, 138
cafer
Pedetes, 128
californica
Aplodontia, 103
californicus
Dipodomys, 111
Lynx, 165
Scapanus, 140
callida
Dasyprocta, 130
Callosciurus
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
Canis
beothucus, 149
familiaris, 150
lupus, 149, 150
Cansumys
canus, 117
canus
Cansumys, 117
capensis
Felis, 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
carissima
Myotis, 146
Carnivora, 149
carolinensis
Sciurus, 106
carpenteri
Hylobates, 136
carrorum
Oryzomys, 123
cascadensis
Lepus, 134
castaneoventris
Sciurus, 103
castanonotus
Sciurus, 99
castanotus
Sciurus, 99
castanops
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
centurto
Ametrida, 143
Cerdocyon
aquilus, 150
germanus, 150
thous, 150
Cervaria
fasciatus, 155
oculeus, 155
Cervidae, 148
Cetartiodactyla, 97
Chaerophon
leucostigma, 145
Cheiromys
laniger, 135
madagascariensis, 135
cherriei
Zygodontomys, 128
Chilonycteris
parnellii, 143 |
pusillus, 143 |
torrei, 143 )
Chionomys |
nivalis, 112
chionopaes
Dicrostonyx, 113
chiriquensis
Guerlinguetus, 106
Sciurus, 106
Chiroptera, 140
Chlorotalpa
tropicalis, 101
chrotorrhinus
Microtus, 114
Chrysochloridae, 101
Chrysochloris
stuhlmanni, 101
tropicalis, 101
chrysogaster
Lemmus, 114
cicognanii
Mustela, 152
cicur
Caluromys, 99
Philander, 99
Cimolesta, 156
cinereoargenteus
Urocyon, 150
cinereus
Sorex, 139
Cingulata, 101
Citellus
obscurus, 107
siccus, 1O7
Claviglis
collaris, 129
soleatus, 129
clavium
Odocoileus, 149
Clethrionomys
gapperi, 113
proteus, 113
collaris
Claviglis, 129
colombiana
Dasyprocta, 130
colonus
Geomys, 110
coloratus
Oryzomys, 123
columbiana
Aplodontia, 103
columbianus
Geocapromys, 132
compacta
Blarina, 137
concolor
Felis, 156
Puma, 156
concreta
Beluga, 148
condylurus
Mops, 145
confucianus
Niviventer, 119
consobrinus
Sciurotamias, 106
cooperi
Synaptomys, 116
Tamias, 109
coryt
Felis, 156
Puma, 156
Corynorhinus
phyllotis, 146
costaricensis
Felis, 156
Puma, 156
cowant
Microgale, 102
Craseomys
aquilus, 113
Crassus
Phenacomys, 116
Cratogeomys
castanops, 111
rubellus, 111
creper
Reithrodontomys, 127
Cricetinae, 117
Cricetomyinae, 117
cricetulus
Saccostomus, 117
crinitus
Peromyscus, 125
Crocidura
bicolor, 137
fuscomurina, 137
geata, 138
hildegardeae, 138
maurisca, 138
phaios, 138
tephragaster, 137
crusnigrum
Pecari, 147
Tayassu, 147
Cryptogale
australis, 101
Cryptomys
hottentotus, 129
TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden
occlusus, 129
whytei, 129
Cryptotis
avia, 138
thomasi, 138
cubanus
Geocapromys, 132
Solenodon, 137
cumberlandius
Geomys, 110
cuppes
Ochotona, 133
curtus
Cebus, 136
Hipposideros, 142
Damaliscus
phillipsi, 149
pygargus, 149
daphaenodon
Sorex, 139
darlingtoni
Eptesicus, 145
Pipistrellus, 145
Dasogale
fontoynonti, 103
Dasymys
alleni, 118
incomtus, 118
Dasypodidae, 101
Dasyprocta
callida, 130
colombiana, 130
leporina, 130
noblei, 130
nuchalis, 130
punctata, 130
Dasyproctidae, 130
Dasypus
hoplites, 101
novemcinctus, 1O1
Dasyuridae, 100
Dasyuromorphia, 100
Daubentonia
madagascariensis, 135
Daubentoniidae, 135
davidanus
Sciurotamias, 106
decaryi
Microgale, 102
declivis
Beluga, 148
degener
Lutra, 152
delectorum
Praomys, 119
deletrix
Vulpes, 150
Delphinapterus
leucas, 148
Delphinus
melas, 148
demidovii
Galago, 135
Dendromurinae, 117
desmarestianus
Heteromys, 111
devius
Oryzomys, 123
diadema
Propithecus, 135
Dicrostonyx
chionopaes, 113
exsul, 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
dorcas
Damaliscus, 149
dorsalis
Tamias, 109
dorsatum
Erethizon, 130
Dremomys
flavior, 104
pernyi, 104
senex, 104
drouhardi
Microgale, 102
douglasii
Tamiasciurus, 109
dupreanum
Eidolon, 98
Echimyidae, 131
Echymipera
australis, 101
kalabu, 100
rufescens, 101
Eidolon
dupreanum, 98
sakalava, 98
elegans
Spermophilus, 108
elongata
Mephitis, 164
elucus
Procyon, 151
Emballonuridae, 142
energumenos
Mustela, 153
Putorius, 153
eEnixus
Microtus, 115
Eonycteris
glandifera, 140
spelaea, 140
Eothenomys
aurora, 113
eva, 113
melanogaster, 113
mucronatus, 113
Epimys
zappeyi, 119
Eptesicus
darlingtoni, 145
phasma, 146
pumilus, 145
eremicus
Hesperomys, 98
Peromyscus, 98, 126
Erethizon
dorsatum, 130
picinus, 130
Erethizontidae, 130
erigens
Hipposideros, 142
Erioryzomys
monochromos, 128
erminea
Mustela, 152, 153
Erophylla
sezekorni, 144
syops, 144
erythraeus
Callosciurus, 103
Euarctos
americanus, 151
sornborgeri, 151
Eulipotyphla, 97
Eupleres
goudotii, 155
major, 155
Eutamias
minimus, 109
neglectus, 109
euxinus
Apodemus, 117
eva
Eothenomys, 113
Evotomys
proteus, 113
exsputus
Sigmodon, 127
173
174 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
exsul
Dicrostonyx, 113
extimus
Sciurus, 106
familiaris
Canis, 150
fasciatus
Cervaria, 155
Lynx, 155
fatuus
Synaptomys, 116
Felidae, 155
Felis
bangsi, 156
capensis, 155
concolor, 156
coryi, 156
costaricensis, 156
floridana, 156
improcera, 156
phillipsi, 155
serval, 155
femurvillosum
Artibeus, 144
ferrugineus
Callosciurus, 103
fervidus
Sigmodon, 128
Fiber
aquilonius, 116
obscurus, 116
rivalicius, 116
zibethicus, 116
fischeri
Haplonycteris, 141
flavicans
Oecomys, 122
Oryzomys, 122
flavidus
Isthmomys, 121
Megadontomys, 121
flavior
Dremomys, 104
flavipes
Tatera, 117
flaviventer
Arctomys, 105
flaviventris
Marmota, 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
fulvescens
Oligoryzomys, 122
fulviventer
Marmosa, 99
fumatus
Myomys, 119
Praomys, 119
Funisciurus
akka, 104
pyrrhopus, 104
victoriae, 104
furculus
Triaenops, 143
furvus
Sigmodon, 128
Urocyon, 150
fuscipes
Neotoma, 98
fuscomurina
Crocidura, 137
fusus
Peromyscus, 126
Galago
demidovii, 135
orinus, 135
Galagoides
orinus, 135
Galagonidae, 135
Galidictis
grandidiensis, 155
grandidieri, 155
gappert
Clethrionomys, 113
geata
Crocidura, 138
Myosorex, 138
Geocapromys
abaconis, 132
columbianus, 132
cubanus, 132
ingrahami, 132
irrectus, 132
Geogale
aurita, 101
orientalis, 101
Geomyidae, 110
Geomys
austrinus, 110
colonus, 110
cumberlandius, 110
floridanus, 110
coffi, 110
pinetis, 110
tuza, 110
Gerbillinae, 117
gerbillus
Leggada, 119
germanus
Cerdocyon, 150
gibbsi
Neurotrichus, 140
giganteus
Pteropus, 141
gigas
Lynx, 155
glaber
Heterocephalus, 129
landifera
Eonycteris, 140
Glaucomys
bangsi, 104
lascivus, 104
makkovikensis, 104
querceti, 105
sabrinus, 104
volans, 104, 105
Glossophaga
longirostris, 144
gloveralleni
Procyon, 151
Sorex, 138
gofft
Geomys, 110
gorgonae
Proechimys, 131
gorilla
Gorilla, 136
Troglodytes, 136
(ey
fo)
Gorilla
gorilla, 136
gossypinus
Hesperomys, 123
Peromyscus, 123, 124,
125
goudotii
: Eupleres, 155
Grammomys
macmillani, 118
granatensts
Sciurus, 106, 107
grandidiensis
Galidictis, 155
grandidieri
Galidictis, 155
grandis
Orthogeomys, 111
Graphiurus
argenteus, 129
griseus, 129
hueti, 129
lorraineus, 129
microtis, 129
murinus, 129
schwabi, 129
surdus, 129
griseus
Graphiurus, 129
griswoldi
Tana, 134
Guerlinguetus
aestuans, 106
chiriquensis, 106
guyannensis
Proechimys, 131
gymnicus
Sciurus, 109
Tamiasciurus, 109
haemobaphes
Callosciurus, 103
Sciurus, 103
Haplonycteris
fischeri, 141
hardyi
Zapus, 112
harpia
Harpiocephalus, 146
Harpiocephalus
harpia, 146
rufulus, 146
harringtoni
Taterillus, 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
hirsutus
Sigmodon, 128
hispida
Suillomeles, 100
hispidus
Sigmodon, 127, 128
Hominidae, 138
hoplites
Dasypus, 101
hottentotus
Cryptomys, 129
hudsonica
Lutra, 152
TYPE SPECIMENS OF RECENT MAMMALS ¢ Helgen and McFadden 175
hudsonicus
Sciurus, 109
Tamiasciurus, 109
Zapus, 112
hueti
Aethoglis, 129
Graphiurus, 129
humulis
Reithrodontomys, 127
hyacinthus
Neurotrichus, 140
hyleae
Proechimys, 131
Hylobates
carpenteri, 136
lar, 136
Hylobatidae, 136
Hylomyscus
alleni, 118
simus, 118
Hylopetes
alboniger, 105
anchises, 105
orinus, 105
phayrei, 105
hypoxanthus
Oenomys, 119
Hypudaeus
nivicola, 112
Ictidomys
badius, 108
tridecemlineatus, 108
Idionycteris
phyllotis, 146
ignava
Marmota, 105
ignavus
Arctomys, 105
ignifer
Rhinolophus, 143
illectus
Oecomys, 122
Oryzomys, 122
impiger
Reithrodontomys, 127
improcera
Felis, 156
Puma, 156
incitatus
Lepus, 134
Sylvilagus, 134
Tapeti, 134
incomtus
Dasymys, 118
Indridae, 135
ingrahami
Geocapromys, 132
innuitus
Mictomys, 116
Synaptomys, 116
insulanus
Peromyscus, 125
insularis
Pteropus, 98
irrectus
Geocapromys, 132
irroratus
Liomys, 112
Isthmomys
flavidus, 121
kalabu
Echymipera, 100
kempi
Tatera, 117
kivuensis
Thamnomys, 120
kodiacensis
Spermophilus, 99
koreni
Microtus, 115
Sorex, 138
labilis
Diplomys, 131
Loncheres, 131
lacustris
Otomys, 120
ladas
Zapus, 112
laetus
Aethosciurus, 106
Lagomorpha, 133
lambertoni
Nesomys, 120
lanatus
Caluromys, 99
laniger
Cheiromys, 135
Pteropus, 96
lanigera
Pteropus, 98
lar
Hylobates, 136
lascivus
Glaucomys, 104
Sciuropterus, 104
Lasiurus
borealis, 146
brachyotis, 146
latimanus
Scapanus, 140
laxatina
Lontra, 142
lecontii
Reithrodontomys, 127
Leggada
ablutus, 118
bufo, 118
gerbillus, 119
Lemmus
chrysogaster, 114
paulus, 114
sibiricus, 114
lepida
Neotoma, 121
Leporidae, 143
leporina
Dasyprocta, 130
Leptailurus
phillipsi, 155
serval, 155
Lepus
alacer, 134
alleni, 133
americanus, 133, 134
arcticus, 133
bairdi, 133, 134
bangsii, 133
cascadensis, 134
incitatus, 134
palitans, 133
paludicola, 134
struthopus, 133
sylvaticus, 134
transitionalis, 134
leucas
Delphinapterus, 148
leucogaster
Scotophilus, 147
leucopus
Hesperomys, 124
Peromyscus, 126
leucostigma
Chaerophon, 145
Mops, 145
levipes
Melomys, 118
Liomys
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
Lontra
canadensis, 152
laxatina, 152
loquax
Sciurus, 109
Tamiasciurus, 109
lorraineus
Graphiurus, 129
lotor
Procyon, 151
louisianae
Odocoileus, 148
lucifugus
Myotis, 146
luctuosa
Alouatta, 135
ludovicianus
Sciurus, 106
lupus
Canis, 149, 150
lutensis
Lutreola, 153
Mustela, 153
Putorius, 153
Lutra
degener, 151
hudsonica, 151
vaga, 151
Lutreola
lutensis, 153
vulgivagus, 154
Lynx
californicus, 155
canadensis, 155, 156
fasciatus, 155
gigas, 155
oculeus, 155
rufus, 155
subsolanus, 156
mabirae
Phataginus, 156
macmillani
Grammomys, 118
Macrogeomys
cavator, 110
pansa, 110
macropygmaeus
Sorex, 138
macrotis
Nycteris, 142
Peropteryx, 142
Macrotolagus
alleni, 133
palitans, 133
macroura
Odocoileus, 148
macrurus
Sorex, 139
madagascariensis
Cheiromys, 135
Daubentonia, 135
Nycteris, 142
Rousettus, 141
major
Eupleres, 155
Vampyrodes, 145
makkovikensis
176
Glaucomys, 104
Sciuropterus, 104
maniculatus
Peromyscus, 98, 124,
125
Manidae, 156
marcanot
Antillogale, 137
Solenodon, 137
Marmosa
fulviventer, 99
mitis, 99
robinsoni, 99, 100
Marmosidae, 99
Marmota
avara, 105
flaviventris, 105
ignava, 105
monax, 105
marsupialis
Didelphis, 100
Martes
americana, 152
atrata, 152
brumalis, 152
maurisca
Crocidura, 138
mayeri
Antechinus, 100
maynardi
Procyon, 151
mearnsi
Saccostomus, 117
medialis
Callosciurus, 103
medioximus
Mictomys, 116
Synaptomys, 116
Megadontomys
flavidus, 121
megaphyllus
Rhinolophus, 143
melanogaster
Eothenomys, TELS}
melanops
Taterillus, 117
melanotis
Oryzomys, 123
melanotus
Praomys, 119
melas
Delphinus, 148
Myoictis, 100
Melomys
alleni, 118
levipes, 118
mollis, 118
moncktoni, 118
rubex, 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
Mesocapromys
nanus, 132
mesomelas
Mephitis, 154
mexicana
Neotoma, 122
mexicanus
Peromyscus, 124
Microgale
brevicaudata, 102
cowani, 102
decaryi, 102
drouhardi, 102
longicaudata, 102
parvula, 102
principula, 102
prolixicaudata, 102
pulla, 102
Microsciurus
alfari, 105
browni, 105
microtis
Graphiurus, 129
Microtus
acadicus, 115
aurora, 113
breweri, 114
chrotorrhinus, 114
enixus, 115
fontigenus, 115
koreni, 115
mucronatus, 113
oeconomus, 115
pennsylvanicus, 114,
115
provectus, 115
ravus, 114
shattucki, 115
terraenovae, 114
micrus
Nesophontes, 136
Mictomys
innuitus, 116
medioximus, 116
MiNntMus
Tamias, 99, 109
minor
Ametrida, 143
Suncus, 139
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
minusculus
Scapanus, 140
miscix
Sorex, 139
misim
Antechinus, 100
mitis
Marmosa, 99
mollipilosus
Tamiasciurus, 109
mollis
Melomys, 118
Stenomys, 120
Molossidae, 145
monax
Marmota, 105
moncktoni
Melomys, 118
monochromos
Erioryzomys, 128
Oryzomys, 125
Thomasomys, 128
Monodontidae, 148
monoensis
Pteropus, 141
Mops
angolensis, 145
condylurus, 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
MUPICUS
Arctogale, 153
Mustela, 153
Putorius, 153
Muridae, 112
Murinae, 117
MUrinUs
Graphiurus, 129
Pseudohydromys, 120
Mus
bactrianus, 119
bairdii, 98
bufo, 118
musculus, 119
tantillus, 119
tenellus, 119
musculus
Mus, 119
Mustela
atrata, 152
brumalis, 152
cicognanii, 152
energumenos, 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
melas, 100
wallacei, 100
wavicus, 100
Myomys
fumatus, 119
Myopus
schisticolor, 115
thayeri, 115
Myosorex
geata, 138
Myotis
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
naso
Antechinus, 100
Rhynchiscus, 142
natator
Oryzomys, 123
navus
Oryzomys, 122
nebrascensis
Hesperomys, 124
Peromyscus, 124
Nectomys
amazonicus, 121
squamipes, 121
neglectus
Tamias, 109
Neomeris
asiaeorientalis, 148
neomexicana
Mustela, 152
neomexicanus
Putorius, 152
Neophocaena
asiaorientalis, 148
phocaenoides, 148
Neosorex
acadicus, 138
palustris, 138
Neotoma
abbreviata, 121
bella, 121
distincta, 122
floridana, 122
fuscipes, 98
lepida, 121
mexicana, 122
rubida, 122
nesaeus
Sciurus, 107
Nesomyinae, 120
Nesomys
lambertoni, 120
Nesophontes
micrus, 136
Nesophontidae, 136
Neurotrichus
gibbsi, 140
hyacinthinus, 140
niger
Sciurus, 106
nigriculus
Peromyscus, 125
niobe
Stenomys, 120
nitellinus
Nyctomys, 122
nivalis
Chionomys, 112
Mustela, 153
nivicola
Hypudaeus, 112
Niviventer
confucianus, 119
noblei
Dasyprocta, 130
notius
Pipistrellus, 147
Putorius, 153
Scabrifer, 147
noveboracensis
Mustela, 153
Putorius, 153
novemcinctus
Dasypus, 101
nuchalis
Dasyprocta, 130
nugax
Myotis, 146
nyasae
Steatomys, 117
Nycteridae, 142
Nycteris
macrotis, 142
madagascariensis, 142
nana, 142
revoili, 97
tristis, 142
Nycticeius
africanus, 147
albiventer, 147
schlieffeni, 147
Nyctomys
nitellinus, 122
sumichrasti, 122
obscurus
Citellus, 107
Fiber, 115
Ondatra, 115
obsoletus
Spermophilus, 99
occidentalis
Paramicrogale, 102
occisor
Mustela, 153
Putorius, 153
occlusus
Cryptomys, 129
Ochotona
cupppes, 133
princeps, 133
saxatilis, 133
Ochotonidae, 133
ochraceus
Thamnomys, 118
Octodontidae, 130
Octomys
barrerae, 130
oculeus
Cervaria, 155
Lynx, 155
ocythous
Urocyon, 150
Odocoileus
americanus, 148
borealis, 148
clavium, 149
louisianae, 146
macroura, 146
osceola, 146
virginianus, 146, 147
Oecomys
bicolor, 122
flavicans, 122
illectus, 122
trabeatus, 122
oeconomus
TYPE SPECIMENS OF RECENT MAMMALS ® Helgen and McFadden 1
Microtus, 115
Oenomys
hypoxanthus, 119
talangae, 119
Oligoryzomys
fulvescens, 122
vegetus, 125
Ondatra
aquilonius, 116
obscurus, 116
rivalicius, 116
zibethicus, 116
orarius
Sciurus, 109
Zapus, 112
ordii
Dipodomys, 111
Dipodops, 111
oreas
Peromyscus, 126
oribasus
Arctogale, 153
Mustela, 153
Putorius, 153
orientalis
Geogale, 101
orientis
Mops, 145
orinus
Galago, 135
Galagoides, 135
Hylopetes, 105
Pteromys, 105
Orthogeomys
cavator, 110
grandis, 111
pansa, 110
pluto, 111
Oryzomys
carrorum, 123
coloratus, 123
devius, 123
flavicans, 122
fulvescens, 122
illectus, 122
melanotis, 123
monochromos, 128
natator, 123
navus, 122
palustris, 123
rostratus, 123
vegetus, 123
osceola
Cariacus, 148
Odocoileus, 146
Otomops
papuensis, 145
Otomyinae, 120
Otomys
anchietae, 120
barbouri, 121
~l
~
lacustris, 120
typus, 121
uzungwensis, 121
owent
Praomys, 119
paca
Agouti, 130
Pachycyon
robustus, 150
paitana
Tupaia, 134
palitans
Lepus, 133
Macrotolagus, 133
palliata
Alouatta, 135
pallidulus
Dipodomys, 111
pallidus
Spermophilus, 99
Tamias, 99
palmarius
Peromyscus, 125
palmarum
Artibeus, 144
palmeri
Dipodomys, 111
Dipodops, 111
paludicola
Lepus, 134
Sylvilagus, 134
palustris
Neosorex, 138
Oryzomys, 123
Sorex, 138
Sylvilagus, 134
pansa
Macrogeomys, 110
Orthogeomys, 110
Pappogeomys
castanops, 111
rubellus, 111
papuensis
Otomops, 145
Paracapromys, 132
Paramicrogale
occidentalis, 102
Paraxerus
byatti, 106
vexillarius, 106
parnellii
Chilonycteris, 143
Pteronotus, 143
parryi
Spermophilus, 99
particeps
Didelphis, 100
parvula
Microgale, 102
paulus
Lemmus, 114
178 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Pecari
crusnigrum, 147
tajacu, 147, 148
torvus, 148
Pedetes
cafer, 125
capensis, 128
taborae, 128
Pedetidae, 128
pennsylvanica
Vulpes, 150, 151
pennsylvanicus
Microtus, 114, 115
Peramelia, 100
pernyt
Dremomys, 104
Perognathus
bangsi, 112
longimembris, 112
Peromyscus
abietorum, 125
ammodytes, 125
anastasae, 124, 125
arenarius, 126
argentatus, 125
attwateri, 124
bairdii, 98
baliolus, 126
bellus, 124
borealis, 124
cacabatus, 124
canadensis, 125
crinitus, 125
eremicus, 98, 126
fusus, 126
gossypinus, 2S 2Ae
125
insulanus, 125
leucopus, 126
maniculatus, 98, 124,
125
mexicanus, 124
nebrascensis, 124
nigriculus, 125
oreas, 126
palmarius, 125
phasma, 126
polionotus, 126
rhoadsi, 126
saturatus, 126
scitulus, 125
subgriseus, 126
texanus, 126
Peropteryx
canina, 142
macrotis, 142
phaea, 142
Peroryctidae, 100
perriert
Propithecus, 135
personatus
Sorex, 139
phaea
Peropteryx, 142
phaios
Crocidura, 138
phasma
Eptesicus, 147
Peromyscus, 126
Pipistrellus, 147
Phataginus
mabirae, 156
tricuspis, 156
phayrei
Hylopetes, 105
Pteromys, 105
Phenacomys
celatus, 116
crassus, 116
ungava, 116
Philander
cicur, 99
philippinensis
Rhinolophus, 143
phillipsi
Damaliscus, 149
Felis, 155
Leptailurus, 155
phocaenoides
Neophocaena, 148
Phocoenidae, 148
Phyllostomidae, 143
phyllotis
Corynorhinus, 146
Idionycteris, 146
picinus
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
auritus, 147
sacrimontis, 147
pluto
Orthogeomys, 111
poeyanus
Solenodon, 137
polionotus
Peromyscus, 126
Praomys
delectorum, 119
fumatus, 119
melanotus, 119
oweni, 119
tullbergi, 119
pratensis
Steatomys, 117
Primates, 135
primus
Callosciurus, 103
princeps
Ochotona, 133
principula
Microgale, 102
priscus
Rhynchiscus, 142
Procyon
elucus, 151
gloveralleni, 151
lotor, 151
maynardi, 151
Procyonidae, 151
Proechimys
burrus, 131
cayennensis, 131
gorgonae, 131
guyannensis, 131
hyleae, 131
semispinosus, 131
prolixicaudata
Microgale, 102
Propithecus
diadema, 135
perrieri, 135
proteus
Clethrionomys, 113
Evotomys, 113
provectus
Microtus, 115
Pseudohydromys
murinus, 120
Pteromys
alboniger, 105
anchises, 105
orinus, 105
phayrei, 105
Pteronotus
parnellii, 143
pusillus, 143
quadridens, 143
Pteropodidae, 140
Pteropus
anetianus, 141
aorensis, 141
ariel, 141
austini, 141
giganteus, 141
insularis, 98
laniger, 98
lanigera, 98
monoensis, 141
rayneri, 141
woodfordi, 141
pulla
Microgale, 102
Puma
bangsi, 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
vulgivagus, 154
xanthogenys, 154
pygargus
Damaliscus, 149
pyrrhopus
Funisciurus, 104
quadridens
Pteronotus, 143
quadrivittatus
Tamias, 99, 109
querceti
Glaucomys, 105
Sciuropterus, 105
rainieri
Aplodontia, 103
Rangifer
arcticus, 149
caboti, 149
caribou, 149
tarandus, 149
terraenovae, 149
ravus
Microtus, 114
raynert
Pteropus, 141
Reithrodontomys
australis, 127
creper, 127
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden 179
humulis, 127
impiger, 127
lecontii, 127
sumichrasti, 127
vulcanius, 127
relictus
Capromys, 131
rendalli
Pipistrellus, 147
repens
Heteromys, 111
revoili
Nycteris, 97
Rhinolophidae, 142
Rhinolophus
alleni, 143
ignifer, 143
megaphyllus, 143
philippinensis, 143
rhoadsi
Peromyscus, 126
Rhynchiscus
naso, 142
priscus, 142
Rhynchonycteris
naso, 142
richardsonii
Mustela, 152
riparia
Arvicola, 114
rivalicius
Fiber, 116
Ondatra, 116
rixosa
Mustela, 153
rixOSUS
Putorius, 153
robinsoni
Marmosa, 99, 100
roboratus
Sorex, 139
robustus
Pachycyon, 150
Rodentia, 103
rostratus
Oryzomys, 123
Rousettus
madagascariensis, 141
rubellus
Cratogeomys, 111
Pappogeomys, 111
rubex
Melomys, 118
rubida
Neotoma, 122
rubricatus
Vulpes, 151
rubricosa
Vulpes, 150, 151
rufa
Aplodontia, 103
rufescens
Echymipera, 101
rufidorsum
Arvicola, 114
rufulus
Harpiocephalus, 145
rufus
Lynx, 155
sabrinus
Glaucomys, 104
Sciuropterus, 104
Saccostomus
cricetulus, 117
mearnsi, 117
sacrimontis
Plecotus, 147
sakalava
Eidolon, 98
saltuensis
Sciurus, 107
sanctaemartae
Sigmodon, 128
sanguinidens
Sorex, 139
saturatus
Peromyscus, 126
saxatilis
Ochotona, 133
Scabrifer
notius, 147
Scalops
aereus, 140
anastasae, 140
texanus, 140
Scalopus
aereus, 140
anastasae, 140
aquaticus, 140
Scandentia, 134
Scapanus
californicus, 140
latimanus, 140
minusculus, 140
schisticolor
Myopus, 115
schlieffeni
Nycticeius, 147
schwabi
Graphiurus, 129
scitulus
Peromyscus, 125
Sciuridae, 103
Sciuropterus
alpinus, 104
bangsi, 104
lascivus, 104
makkovikensis, 104
querceti, 105
sabrinus, 104
silus, 104
volans, 104, 105
Sciurotamias
consobrinus, 106
davidianus, 106
thayeri, 106
Sciurus
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
morulus, 107
nesaeus, LOT
niger, 106
orarius, 109
saltuensis, 107
variabilis, 107
vicinus, 106
vulpinus, 106
yucatanensis, 106
Scotinomys
apricus, 127
teguina, 127
xerampelinus, IAG
Scotophilus
altilis, 147
leucogaster, 147
scrutator
Mephitis, 154
semispinosus
Proechimys, 131
senex
Dremomys, 104
seorsus
Zygodontomys, 128
serval
Felis, 155
Leptailurus, 155
Setifer
setosus, 103
setosus
Setifer, 103
sezekorni
Erophylla, 144
shattucki
Microtus, 115
sibiricus
Lemmus, 114
SICCUS
Citellus, 107
Sigmodon
austerulus, 127
borucae, 127
exsputus, 127
fervidus, 128
furvus, 128
hirsutus, 128
hispidus, 127, 128
sanctaemartae, 128
spadicipygus, 128
Sigmodontinae, 121
silus
Sciuropterus, 104
simus
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
macropygmaeus, 138
macrurus, 139
miscix, 139
palustris, 138
personatus, 139
roboratus, 139
sanguinidens, 139
tundrensis, 138
ultimus, 138
vir, 139
Soricidae, 137
sornborgeri
Euarctos, 151
Ursus, 151
sOoror
Tatera, 117
spadicipygus
Sigmodon, 128
spelaea
Eonycteris, 140
Spermophilus
alashanicus, 107
armatus, 107
badius, 108
elegans, 108
kodiacensis, 99
obsoletus, 99
pallidus, 99
parryi, 99
180 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
spilosoma, 99
texensis, 108
tridecemlineatus, 99,
108
Spilogale
ambarvalis, 155
putorius, 155
spissigrada
Mephitis, 154
squamipes
Nectomys, 121
Steatomys
nyasae, 117
pratensis, 117
Stenomys
mollis, 120
niobe, 120
stevensi, 120
verecundus, 120
stevensi
Melomys, 118
Stenomys, 120
striatus
Tamias, 107
struthopus
Lepus, 133
stuhlmanni
Chrysochloris, 101
stygius
Heterocephalus, 129
subgriseus
Peromyscus, 126
subsolanus
Lynx, 156
Suillomeles
hispida, 100
sumichrasti
Nyctomys, 122
Reithrodontomys, 127
Suncus
ater, 139
minor, 139
varilla, 139
surdus
Graphiurus, 129
sylvaticus
Lepus, 134
Sylvilagus
alacer, 134
brasiliensis, 134
floridanus, 134
incitatus, 134
paludicola, 134
palustris, 134
transitionalis, 134
Synaptomys
~ borealis, 116
cooperi, 116
fatuus, 116
innuitus, 116
medioximus, 116
Syntheosciurus
brochus, 108
syops
Erophylla, 144
taborae
Pedetes, 128
tajacu
Pecari, 147, 148
talangae
Oenomys, 119
Talpidae, 140
Tamias
borealis, 109
cooperi, 108
dorsalis, 108
minimus, 99, 109
neglectus, 109
pallidus, 99
quadrivittatus, 99
striatus, 109
townsendii, 108
venustus, LOY
Tamiasciurus
douglasii, 109
gymnicus, 109
hudsonicus, 109
loquax, 109
mollipilosus, 109
Tana
griswoldi, 134
tana, 134
tana
Tana, 134
Tupaia, 134
tantillus
Mus, 119
Tapeti
incitatus, 134
tarandus
Rangifer, 149
Tatera
flavipes, 117
kempi, 117
soror, 117
valida, 117
Taterillus
harringtoni, 117
melanops, 117
Tayassu
crusnigrum, 147
torvus, 148
Tayassuidae, 147
teguina
Akodon, 127
Scotinomys, 127
tenellus
Mus, 119
Tenrecidae, 101
tephragaster
Crocidura, 137
terraenovae
Arvicola, 114
Microtus, 114
Rangifer, 149
texanus
Peromyscus, 126
Scalops, 140
texensis
Spermophilus, 108
Thamnomys
kivuensis, 120
ochraceus, 118
venustus, 120
thayeri
Myopus, 115
Sciurotamias, 106
thomasi
Cryptotis, 138
Thomasomys
monochromos, 128
Thomomys
atrodorsalis, 111
umbrinus, 111
thous
Cerdocyon, 150
Thyroptera
albigula, 147
albiventer, 147
tricolor, 147
Thyropteridae, 147
torquatus
Dicrostonyx, 113
torrei
Boromys, 131
Chilonycteris, 143
torvus
Pecari, 148
Tayassu, 148
townsendii
Tamias, 109
trabeatus
Alouatta, 135
Oecomys, 122
transitionalis
Lepus, 134
Sylvilagus, 134
Triaenops
aurita, 143
furculus, 143
tricolor
Thyroptera, 147
tricuspis
Phataginus, 156
tridecemlineatus
Spermophilus, 99, 108
trinotatus
Zapus, 112
tristis
Nycteris, 142
Troglodytes
gorilla, 136
tropicalis
Chlorotalpa, 101
Chrysochloris, 101
tullbergi
Praomys, 119
tundrensis
Sorex, 138
Tupaia
paitana, 134
tana, 134
Tupaiidae, 134
turpis
Hipposideros, 143
tuza
Geomys, 110
Tympanoctomys
barrerae, 130
typus
Otomys, 121
ultimus
Sorex, 138
umbratus
Platyrrhinus, 144
Vampyrops, 144
umbrinus
Thomomys, 111
ungava
Phenacomys, 116
Urocyon
aquilus, 150
cinereoargenteus, 150
furvus, 150
ocythous, 150
Ursidae, 151
Ursus
americanus, 151
sornborgeri, 151
UZUNZWENSIS
Otomys, 121
vafra
Vulpes, 150, 151
vaga
Lutra, 152
valida
Tatera, 117
Vampyrodes
caraccioli, 145
major, 145
Vampyrops
umbratus, 144
zarhinus, 144
variabilis
Sciurus, 107
varilla
Suncus, 139
vegetus
Oligoryzomys, 123
Oryzomys, 123
venustus
Tamias, 109
Thamnomys, 120
verecundus
Stenomys, 120
Vespertilionidae, 145
vexillarius
Paraxerus, 106
vicinus
Sciurus, 106
victoriae
Funisciurus, 104
vir
Sorex, 139
virgatus
Agouti, 130
virginiana
Didelphis, 100
virginianus
Odocoileus, 148, 149
vison
Mustela, 153, 154
Putorius, 153
Viverridae, 155
volans
Glaucomys, 104, 105
Sciuropterus, 104, 105
vulcanius
Reithrodontomys, 127
vulgivaga
Mustela, 154
vulgivagus
Lutreola, 154
Putorius, 154
vulpes
Vulpes, 150, 151
Vulpes
bangsi, 151
deletrix, 150
pennsylvanica, 150, 151
rubricatus, 151
rubricosa, 150, 151
vafra, 150, 151
vulpes, 150, 151
vulpinus
Sciurus, 106
wallacei
Myoictis, 100
wavicus
Myoictis, 100
whytei
Cryptomys, 129
woodfordi
Pteropus, 141
xanthogenys
Putorius, 154
xerampelinus
Akodon, 127
Scotinomys, 127
yucatanensis
Sciurus, 106
TYPE SPECIMENS OF RECENT MAMMALS * Helgen and McFadden
zappeyt
Epimys, 119
Zapus
acadicus, 112
hardyi, 112
hudsonicus, 112
ladas, 112
orarius, 112
trinotatus, 112
zarhinus
Vampyrops, 144
zibethicus
Fiber, 116
Ondatra, 116
Zygodontomys
brevicauda, 128
cherriei, 128
seorsus, 128
181
oe j ee SM 4
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é
US ISSN 0027-4100) _
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
VOLUME 157, NUMBER 3
AMBRIDGE, MASSACHUSETTS, U.S.A. 14 November 2001
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
OR DISTRIBUTED BY THE
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OccASIONAL PAPERS ON Mo tusks, 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 Emest 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
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© 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. CADLE'
CONTENTS
ENTERS Sa OPT ee tay ee 183
[Reganrrayeran’ 2 oki eicteie el ae ee es 183
Mien @ GLU CHOI yen tek PLUM Pes REE TUE a Re 184
Materials and Methods 000 184
Description of a New Species of Stenocersus 187
Stenocercus prionotus new species —.... 187
Diagnosis and Comparisons 2 191
BBY SS CIs tol Opes ee eee ee as sees oe 193
Distribution Patterns in Stenocercus prionotus
ces OE EE aed ae ad oe Lie ee 197
Patterns of Sympatry and Geographic Variation
in Stenocercus prionotus, and the Need for
Additional Fieldwork 200
Natural History of Stenocercus prionotus 205
Comparison of Stenocercus prionotus with S.
COCULCUS Se eke MERE A Ni ee 2 ee Nea RE 206
Distributions of Stenocercus prionotus and S.
caducus in Eastern Bolivia ___.................. 210
Is the Distribution of Stenocercus aculeatus
GHGHTONIVGI Ci a5 Reet pee 270s Meee cee ee ee 212
Key to Species of the “Ophryoessoides group”
Git SHOMOCEROUS Did}
Ac noyylecearncints: 3 ae ee 215
Mppendix: Specimens Examined 0). 216
ihiterature Cited 228. 2 218
ABSTRACT. A new species of iguanid lizard, Steno-
cercus prionotus, is described Gorn eastern Peru and
Bolivia (known range from 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, iiliots 60513.
Research Associate, Department of Herpetology,
Museum of Comparative Zoology.
Bull. Mus. Comp. Zool.,
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 observed in ct 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 than 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 Pert Orien-
tal y ob Bolivia. Se conoce la nueva especie desde el
departamento de San Martin, Pert, 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 l6bulo
escamoso tinico que oculta una porcidén 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
157(3): 183-222, November, 2001 183
154
Argentina y del Paraguay. Stenocercus prionotus se
distingue de S. caducus 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 Pert tienen una cresta vertebral mas alta que la
cresta en poblaciones del sur de Perti y Bolivia. Las
poblaciones del norte de Pert también son amplia-
mente simpatricas con dos otras especies del “grupo
Ophryoessoides,” S. aculeatus y S. fimbriatus. Sin em-
bargo, en el sur del Peru y Bolivia no se conocen lo-
calidades donde se encuentra S. prionotus simpatrica
con otras especies del grupo. Postulo que la cresta ver-
tebral mas alto en las poblaciones nortefias de S. prion-
otus functiona como un sefial para reconocimiento de
especies en comunidades donde hay varias especies
simpatricas. Se provee una clave para las especies del
“grupo Ophryoessoides” y se resumen las distribu-
ciones de las especies Peruanas y Bolivianas.
INTRODUCTION
Stenocercus sensu Frost (1992), includ-
ing the nominal genera Ophryoessoides and
Proctotretus, isa “moderately diverse assem-
blage of South American iguanid lizards
(eos Macey et al., 1997; Soule 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 Peru, 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 Ophryoessoides ‘before the
current understanding of these names
came into use (see Etheridge, 1966; Frost,
1992). Rodriguez and Gadlet (1990) left the
new species cnameless in a checklist pend-
ing resolution of its status. The new species
is ‘apparently closely related to Stenocercus
caducus (Cope) and is described herein.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
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 Duméril and
Bibron sensu lato (including Ophryoesso-
ides Duméril and Proctotretus Duméril
and Bibron). Externally, the new species is
similar to those species that Fritts (1974)
placed in Ophryoessoides, 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
“Ophryoessoides group” are aculeatus
O'Shaughnessy, 1879; caducus Cope,
1862; dumerilii Steindachner, 1867; ery-
throgaster Hallowell, 1856; fimbriatus Avi-
la-Pires, 1995: huancabambae Cadle,
1991: iridescens Giinther, 1859; limitaris
Cadle, 1998; scapularis Boulenger, 1901;
tricristatus Duméril, 1851; ee two un-
described species noted later in this paper
under Key to Species of the “Ophryoesso-
ides group” of Stenocercus. I use the term
“Ophryoessoides 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.
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
Mable ws
All measurements are in millimeters.
The abbreviation SVL refers to the head—
body length, from snout to vent. The con-
185
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
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186
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
S. prionotus
S. aculeatus
S. fimbriatus
S. scapularis
S. caducus
200 400 600 800 1000km
Figure 1.
Distribution of species of Stenocercus emphasized in this paper (western South America, Ecuador to Paraguay and
northern Argentina). Open symbols for S. fimbriatus 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
vary 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 Grails (1991):
Neck and Body Folds and Crests. In
contrast to many species of Stenocercus,
neck folds are usually absent or weakly de-
soides group’
veloped in the “Ophryoessoides 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-
- an antehumeral crest, which
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
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 “Ophryoessoides 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 fie
neck region, where it is confluent with the
supra-auricular crest. In other species it is
present only anteriorly or posteriorly.
Posthumeral (Axillary) and Postfemoral
Mite Pockets. Type 1—pocket absent; no
skin modification. Type 2—rudimentary
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-
illary flap. This structure is described more
fully later.
Angulate temporal scales are distinctly
enlarged, keeled scales posterior to, and in
line with, the superciliary 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-
187
pesto as used in some literature
(e.g., Avila-Pires, 1995). In several species
of eae 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 LIS loo 2s M9931 997,), 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 fhe on-line
versions of the Peru and Bolivia gazetteers
of the U.S. Board on Geographic Names
at the GEOnet® Names Server: http://
164.214.2.59/ens/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 caducus (Cope, 1862): Boulenger (1898),
specimen from “Barraca, Rio Madidi” [Bolivia]
(BMNH 98.6.9.4) and probably two other northern
Bolivian localities discussed in the text.
Ophryoessoides caducus (Cope, 1862): Fugler (1989:
63), specimens from San Marcos Ranch, El Beni
Department, Bolivia, including ROM 12815.
Ophryoessoides aculeatus (O'Shaughnessy, 1879):
Fugler (1983, 1986, 1989), specimens from Tumi
Chucua, El] Beni Department, Bolivia (USNM par-
atypes).
Ophryoessoides sp.: Rodriguez and Cadle (1990),
specimen from Cocha Cashu, Manu National Park,
Madre de Dios Department, Peru (MCZ 150243).
* Stenocercus prionotus 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 Zoology, Woll, ISS INO. 3
. prionotus
. aculeatus
. fimbriatus
. scapularis
. huancabambae
. limitaris
. Iridescens
me SN
Zul
UN
kilometers
\
al
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 <0.87.
—
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. iridescens continues farther north
in western Ecuador than the area covered by the map.
190
Holotype (Figs. 3-5). United States
National Museum of Natural History
(USNM) 193683 (field number WCS
2421). PERU: Drepro. HuANUCo: Jardin
Botdnico 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.
Paratypes 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 Monz6n, 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 Monz6n, 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 Paratypes. PERU: DEpPpTo.
HuANuco: Buena Vista, Valley of the
Chimchao [= Rio Chinchao] [approximate-
ly (9°35, 7a 52 W)| (l=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] (January 1969, Hans W. Koep-
cke), KU 179058. [DEPToO. LORETO]:
Elastern| 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|arvey] 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
247468-69, 247680; (23 May 1986, Vic-
tor R. Morales), USNM 269022. DEPpTo.
Puno: Prov. Sandia: Tambopata, San Juan
(dell Oro], 1520-m or S000: fe 4co7s:
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,
B.Orm: lO6280%Ss (6125.1 (25 July 1984,
Rainer Schulte), KU 212629.
BOLIVIA: DEpTo. BENI: 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, is Lovisek), ROM 12815. [DEPTo.
La Paz]: Barraca, Rio Madidi [12°35’S,
67°02'W] (1893, Luigi Balzan), BMNH
98.6.9.4.
Distribution (Fig. 1). Stenocercus prion-
otus 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-
em Peru (Fig. 1). See Distribution Pat-
terns in Stenocercus prionotus for further
discussion.
Etymology. The epithet prionotus is de-
3 Hilda Heller’s 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 the Rio Tambopata at 5000 feet.” With some hes-
itancy I identify Heller’s locality as the town known
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
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. caducus.
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 preserved: 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
OM
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-preserved 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. fumbriatus Avila-Pi-
res, 1995: S. huwancabambae Cadle, 1991:
and S. scapularis (Boulenger, 1901). An
undescribed species occurs in the Rio
Maranon valley of eastern Peru (see Key
to Species of the “Ophryoessoides Group”
192
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. prionotus 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 Peru 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. aculeatus), S. prionotus
is distinguished from S. fumbriatus (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 parentheses) in lack-
ing a postfemoral pocket (moderate to
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
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 superciliary 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
“Ophryoessoides 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
cercus except the “Ophryoessoides 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 dimorphism, 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. aculeatus, two other spe-
cies of Stenocercus are known from local-
ities close to or sympatric 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 pr ominently 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); str ‘ongly 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 mod-
erately enlarged, 2 mediocentral scales
much larger than the others. Five or 6
scales across the supraocular area at its
S FROM PERU AND BOLIVIA ¢ Cadle 193
widest part. Interparietal distinct and elon-
gate, diamond-shaped or pentagonal (apex
posteriorly). P Parietal eye Tobie. A pair of
parietals in contact behind the interpari-
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
coales are intercalated at the juncture of
the parietal, postparietal, and/or occipital
(ee) buen 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 pr ojecting vane. Keels of adjacent an-
gulate temporals 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 “4 to % 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 sharply 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
| GBB ERS
a E( =v Ne ce
ER
CS
SEES hen ES
Las es
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; |, interparietal; O, occipital; P, parietal; PP, postparietal.
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
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.
Posthumeral and Postfemoral Mite
Pockets. Posthumeral mite pocket a deep
cavity (Type 4) with a prominent axillary
flap concealing the anteroventral aspect
(Fig. 5). Postfemoral pocket absent (Type
ib)
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
195
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
% to % 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 Proportions. Largest male
(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 % 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 running between the eyes on the
top of the 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 a 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 preservative. Poorly preserved
specimens may be more or less uniform
brown all over, although obscure pattern
elements 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 may 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 very prom-
inent). Flanks usually unicolor and some-
what darker than the dorsum between the
dorsolateral crests; however, some 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
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-
U7
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 Peru. 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-
afion 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 Maranon.
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
TABLE 2
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
PATTERNS OF PRESUMED SYMPATRY OF STENOCERCUS IN EASTERN PERU. ONLY SPECIES OF THE
“OPHRYOESSOIDES GROUP” AND THE SUPERFICIALLY SIMILAR SPECIES, S. ROSEIVENTRIS, ARE LISTED.! 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 i Tingo Cuzco Explorer's
Region W. Loreto? Hermosa Mishollo Maria Manu? Balta Amazonico Inn
S. prionotus x x x x x
S. aculeatus x
S. fimbriatus x x x x x
S. roseiventris x x x x
'' The only other species of Stenocercus known from primarily the lowlands and lower Andean foothills in
this region is S. crassicaudatus, 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. fimbriatus and S. roseiventris are sympatric at Pakitza, whereas S. prionotus is
known only from Cocha Cashu.
lizards obtained by Dixon and Soini (1986)
and additional collections from 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 seems to be absent from these sites.
Documenting and explaining patterns of
absence is always difficult, but the appar-
ent absence of Stenocercus prionotus from
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 more than two
species of the “Ophryoessoides group” of
Stenocercus are sympatric. 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 tat region. Numerous spec-
imens of Stonocerels of at least five spe-
cies (S. boettgeri, S. crassicaudatus, S. for-
mosus, S. scapularis, and S. variabilis*)
* 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-
have been collected along an elevational
cribing these specimens to “Palca, Bolivia” 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
itinerary 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
Palea, 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
chrysopygus from northern Peru.
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle
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 (Hudénuco Department)
and Cocha Cashu in Manu National Park
(Madre de Dios Department), a gap of
some 600 km that includes the entire up-
gg
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 Fieldwork). 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 with 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-
ucus 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 Ghuilsmnann 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].° 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 (Métraux,
200
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” (AMNH 22450, 22532)
from Rurrenabaque, Bolivia [E] Beni De-
partment; 14°28’S, 67°34'W]. These were
reidentified in 1971 by Thomas H. Fritts
as Stenocercus caducus but they are pres-
ently missing from the AMNH collection
(Linda Ford, in litt., February 1999). Rur-
renabaque is very close to Balzan’s locality
(2) above. Based solely on presumed hab-
itats and presently known distributions of
S. prionotus and S. caducus in eastern Bo-
livia (see text; Fig. 1), I suspect that
AMNH 22450 and 22532 are most likely
S. prionotus. They should be reexamined
if they are ever located. As an outside pos-
sibility, any of the unverified specimens
from these Bolivian localities could rep-
resent S. fimbriatus, which is now known
from southern Peru (Appendix).
PATTERNS OF SYMPATRY AND
GEOGRAPHIC VARIATION IN
STENOCERCUS PRIONOTUS, AND THE
NEED FOR ADDITIONAL FIELDWORK
Stenocercus prionotus varies geographi-
(S OO
cally in several characters, most notably in
the height of the vertebral crest. Because
1942). According to Métraux (1942: 15), by the end
of the 19th century when Balzan collected his spec-
imens the decimated population of the Mosetenes
was concentrated in the three “Misiones Mosetenes”
of San Miguel de Muchanes (15°14’S, 67°39'W), San-
ta Ana (15°31'S, 67°30’W), and Covendo (15°49’S,
67°06'W). The approximate coordinates given in the
text are the average for these three sites. According
to several maps with elevational contours indicated,
Muchanes and Covendo are between 500 and 1,000
m and Santa Ana is less than 500 m in elevation.
Mathews (1879) also discusses these missions.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
of the potential role of the crest in either
intra- or interspecific communication it
seems appropriate to discuss patterns of
geographic variation in S. prionotus in the
context of the distribution of other species
of the “Ophryoessoides group.” In this re-
gard, patterns of sympatry among species
of this group in Peru and Bolivia are es-
pecially relevant.
Patterns of Sympatry of Species in the
“Ophryoessoides Group.” The most com-
plex distributional patterns for the
“Ophryoessoides group” are in northern
and central Peru, where six species occur
(Stenocercus aculeatus, S. fimbriatus, S.
prionotus, S. huancabambae, S. scapularis,
and an undescribed species; Fig. 2). In
contrast to the other species, S. huanca-
bambae and the undescribed species occur
in comparatively dry deciduous forests
west of the known distributions for the
other species. Their distributions probably
do not overlap the other species and they
will not be considered further here. Based
on known latitudinal and elevational dis-
tributions, three species (S. aculeatus, S.
fimbriatus, and S. prionotus) are probably
broadly sympatric in northern Peru, al-
though sympatry is documented only for
pairs of these at three localities (Fig. 2;
Table 2). The habitat preferences of the
species in the region of sympatry are un-
known.
Whether Stenocercus scapularis is sym-
patric with S. prionotus is less clear. Sten-
ocercus scapularis is known from two
widely separated areas in Hudnuco and
Puno departments (Peru) (Fig. 1; Appen-
dix), and it seems to be elevationally par-
apatric to S. prionotus in northern Peru
(Hudnuco Department). In this area, S.
scapularis occurs above 1,000 m and S.
prionotus is not known above 700 m. In
Puno Department, S. prionotus is known
from a series obtained by Hilda Heller at
San Juan del Oro (1,520 m elevation),
which elevationally overlaps the distribu-
tion of S. scapularis in the same region
(1,000-1,830 m). However, these two spe-
cies have not been taken at the same lo-
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ¢ Cadle
cality in southern Peru. Interestingly, al-
though the largest series of S. prionotus
available from a single locality (San Juan
del Oro) is in the elevational range of S.
scapularis, no specimens of that species
were obtained by Heller during a month-
long stay at the site.
In southern Peru and Bolivia, Stenocer-
cus prionotus is not known to be sympatric
with other members of the “Ophryoesso-
ides group.” However, based on the oc-
currence of S. prionotus and S. fumbriatus
at two nearby localities within Manu Na-
tional Park (Cocha Cashu and Pakitza,° re-
spectively), sympatry for these two species
is expected along the Andean front in this
region. In southern Peru and Bolivia, S.
prionotus is also broadly sympatric with S.
roseiventris, a large terrestrial species that
is superficially similar to species of the
“Ophryoessoides group.” However, these
species have been taken together at only
one locality, Explorer’s Inn.
To summarize these patterns of sympat-
ry, the distributions of three species of the
“Ophryoessoides group” overlap broadly in
northern Peru. Stenocercus prionotus is
known to be sympatric with at least one
other species of the group in Hudnuco and
Loreto departments. No species of the
“Ophryoessoides group” are known to be
sympatric in southern Peru and Bolivia.
Geographic Variation. Geographic vari-
ation pertains to both quantitative and
qualitative characters in Stenocercus
prionotus. Specimens from the northern
part of the range have higher average
counts of midbody scales than southern
specimens, which is shown graphically in
Figure 8. However, there is broad overlap
in the counts from opposite ends of the
range. A reverse trend (lower counts in the
north), occurs in the number of vertebral
(dorsal crest) scales (Fig. 9); this reflects
the reduced prominence, and hence small-
° The record of Stenocercus fimbriatus from Pak-
itza (Appendix) extends the known range for this spe-
cies south by approximately 200 km ‘from Balta in
Ucayali Department, Peru (Avila-Pires, 1995).
201
er scales, of the crest in southern popula-
tions. None of these differences in scale
counts is statistically significant.
The height of the vetebrl crest shows
strong Glinal variation in Stenocercus
prionotus. Populations from northern and
central Peru have higher crests than those
from southern Peru Sand Bolivia (Fig. 10).
However, the absence of specimens be-
tween Hudnuco Department in central
Peru and Manu National Park (Madre de
Dios Department) in southern Peru (Fig.
1) makes it impossible to analyze this trend
in detail. Specimens from the former re-
gion have a high crest typical of all speci-
mens from that area and farther north,
whereas specimens from Madre de Dios
Department and farther south have dis-
tinctly lower crests. The crest is not
strongly sexually dimorphic in either re-
gion.
Whether the transition in crest height is
abrupt or gradual between central and
southern Peru is unclear. Indeed, it is un-
clear whether intervening populations of
Stenocercus prionotus exist. Both patterns
of clinal variation have been extensively
documented empirically and theoretically
for many organisms (Endler, 1977). The
pattern of geographic variation in S. prion-
otus could conform to any of several in
Endlers (1977: 4) classification scheme.
However, present knowledge of distribu-
tions and character variation is most simi-
lar to Endler’s “differentiated disjunction”
pattern, wherein disjunct populations of a
species diverge in one or more characters.
Further speculation is not fruitful in the
absence of more extensive collections from
the region of disjunction in eastern Peru.
A Hypothesis of Causation for Geo-
graphic Differentiation in the Vertebral
Crest of Stenocercus prionotus. Variation
in the height of the vertebral crest is a
prominent distinction between northern
and southern populations of Stenocercus
prionotus. This variation calls for an expla-
nation while at the same time recognizing
that, in the absence of experimental or
field studies, ascribing causation to pat-
48 e@ @
47 )
46
@
45 @ @
44 ee
43 ee ee
42 eee ee
im)
2
gS 41 e
7p)
>
3 40 @ @®e ©
ie}
3
S 39 ) ) @
38 ee
37 ® @
36 ee
35
2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
6.5- 9.3-
7.2 9.6
11.1-
11.8
12.6-
12.9
25.4-
25.6
20.8-
21.3
17.5-
18.3
14.2
Latitude
Figure 8. Geographic variation in the number of midbody scales in Stenocercus prionotus (@) and S. caducus (Ml). Each
symbol represents one specimen; counts for all specimens examined are plotted. The horizontal axis represents degrees south
latitude for each locality converted to a decimal value (e.g.,
17°30’ converted to 17.5); geographically contiguous localities were
arbitrarily combined to yield the latitudinal ranges given. The S. caducus specimens examined were biased toward Bolivian
specimens.
terns of geographic variation is difficult.
Vertebral crests in iguanids are visual sig-
nals. The crests in sexually dimorphic spe-
cies are involved in intr aspecific behavioral
encounters, especially intrasexual aggres-
sion (e.g., Watkins, 1998). Nonetheless,
despite strong geographic differentiation,
the vertebral crest in Stenocercus priono-
tus is not strongly sexually dimorphic in
any part of its range. Thus, a role in in-
trasexual behavioral signaling seems un-
likely to be the primary function of the
crest in this species. Intersexual signalling
is a possibility, but this would not explain
why northern and southern populations
differ so strongly in this character.
Populations of Stenocercus prionotus
with the highest vertebral crests occur in
the area of highest species density and
documented Supe for the “Ophryoes-
soides group” (Table 2 and above discus-
sion). Not only do northern populations of
S. prionotus have the highest vertebral
crests within this species, but in northern
and central Peru S. prionotus has a much
more prominent vertebral crest than other
species of the “Ophryoessoides group” (S.
aculeatus approaches it most closely). In
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle 203
43
42
A
40
39 e e
38 e e e
37 ee
36 ee e ee
35 e e
34 e e
Vertebral Crest Scales
6.5- 9.3- 11.1- 12.6-
7.2 9.6 11.8 12.9
17.5-
15.3 18.3 21.3 25.6
14.2 14.6- 20.8- 25.4-
Latitude
Figure 9. Geographic variation in the number of vertebral crest scales in Stenocercus prionotus and S. caducus. Symbols and
axes are the same as Figure 8.
contrast, in southern Peru and northern
Bolivia the distribution of S. prionotus
overlaps only S. scapularis and (perhaps)
S. fimbriatus of the “Ophryoessoides
group, but S. prionotus is not known to
be sympatric with either of these species.‘
7 The only case of sympatry of any of these with
other species of Stenocercus of which I am aware
involves S. prionotus and S. roseiventris at Explorer's
Inn (cf. Table 2), but S. roseiventris is phenotypically
dissimilar to species of the “Ophryoessoides group”
in having shorter limbs, different color pattern, spiny
tail, and different body form.
The vertebral crest of S. prionotus in
southern Peru and northern Bolivia is sim-
ilar in development to that in allopatric
populations of S. aculeatus.
I conjecture that the strongly developed
vertebral crests of Stenocercus prionotus
in northern Peru may be related to the
presence of multiple sympatric congeners
in that part of the range, and that the crest
functions as a species recognition charac-
ter. If true, this explanation for variation in
crest height would have no precedent
among lizards. The only analogous situa-
204 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
(a) prionotus AS
Huanuco, Peru
67 mm (es)
(b) prionotus FE
Puno, Peru
73 mm @
(Cc) prionotus
Beni, Bolivia
81 mm ©)
ae or
(d) caducus
Sta. Cruz, Bolivia
72 mm
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
tion seems to be dewlaps in Anolis, in
which species recognition has been pos-
tulated as a biological role for dewlap color
and size in complex assemblages of these
lizards (Williams and Rand, 1977; Losos
and Chu, 1998).
The Need for Additional Fieldwork.
Only field observations and experimental
studies of the function of the vertebral
crest in Stenocercus prionotus can deter-
mine whether the functional hypothesis
advanced here is correct. Ideally, parallel
studies should be conducted in the area of
sympatry with other species of the
“Ophryoessoides group,” as well as in the
area with no known sympatric species of
that group. That design would permit an
evaluation of any context-dependent use of
the crest in these populations. Two sym-
patric species are known from the vicinity
of Tingo Maria (Hudnuco Department;
Table 2), a very accessible area for study.
Two other reasons for additional com-
prehensive fieldwork on these species are
indicated. First, we need documentation
of the extent to which species of the
“Ophryoessoides group” are sympatric or
syntopic. Particular targets should be the
northern part of the range of Stenocercus
prionotus, where S. aculeatus and S. fim-
briatus are known to have overlapping
ranges; and Manu National Park in south-
ern Peru, a reasonably accessible area
where S. fumbriatus and S. prionotus are
known to be closely allopatric (but not
sympatric) on a microgeographic scale (Ta-
ble 2). Second, the character differences
between the northern populations of S.
prionotus and those of southern Peru and
Bolivia could indicate that two or more
species reside within my concept of this
species. Fieldwork concentrated in the
e=
205
geographic gap between Tingo Maria and
vicinity (Hudnuco Department) and Cocha
Cashu (Madre de Dios department), the
two most proximate localities of the north-
ern and southern population groups, is
needed to verify whether intermediate
populations occur. The possibility that
these might be two allopatric species sep-
arated by a broad geographic gap should
not be dismissed wathout further evalua-
tion.
NATURAL HISTORY OF STENOCERCUS
PRIONOTUS
Few natural history observations exist
for Stenocercus prionotus. Wade C. Sher-
brooke (field notes) found the holotype
running in the relatively open floor of a
bamboo garden at about 1200 h. USNM
193685 was taken along a trail within rain-
forest. Brief notes ascociated with the se-
ries from Explorer’s Inn (Madre de Dios
Department, Peru; USNM 247468-69,
247680, 269022) indicate that specimens
were obtained during the day on the
ground from clearings around the lodge,
although USNM 269022, an adult female
(91 mm SVL), was on a leaf 40 cm above
the ground. MCZ 150243 was retrieved
from a mist net near the edge of tall flood-
plain forest at Cocha Gaskin: In the dry
season (September—December) of a low-
land rainforest in the vicinity of Tumi Chu-
cua (Beni Department, Bolivia), Fugler
(1986: table 5) found Stenocercus priono-
tus (reported as Ophryoessoides aculeatus )
in varzea (seasonally inundated rainforest)
but not in terra firme rainforests of the
area; females with enlarged eggs were
found in early November (Fugler, 1986:
table 4).
Hilda H. Heller provided the following
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
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-
dicating 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
tic 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. fumbria-
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,
*T 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
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 improbable 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
similarity 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
prionotus from S. caducus, and the species
'° Im 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, Par aguay, and Argentina are, in fact, all
the same taxon. Considerable variation exists in some
aspects of coloration and scale characters in these
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 * Cadle
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 prionotus 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. prionotus.
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 prionotus from S. caducus in south-
"Tam uncertain how Fugler (1983, 1986, 1989)
distinguished Bolivian 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 listed 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).
bo
0
qq
Figure 11. 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 <8.5.
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
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 from southern Bo-
livia, including a series of 31 specimens
(10 males =60 mm SVL) from the vicinity
of Santa Cruz. A similar 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 tem-
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
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 preserved 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 very sim-
ilar judging from descriptions of S. cadu-
cus in the literature (Gallardo, 1959:
Scrocchi et al., 1985; Cei, 1993).
A Possible Difference 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). Compare to Figure 7.
210
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 Subb=
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 3 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 (N
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 ewaal size dimor-
phism provides another character distin-
guishing S. prionotus and S. caducus. Data
presented in Table 1 suggests that other
species of the “Ophryoessoides group”
may be size dimorphic (S. fimbriatus, S.
huancabambae, and ?S. scapularis) or not
(S. aculeatus), and either males (S. huan-
cabambae) 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
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 aoa specimens
in these reports.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
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 Cruz 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 somewhat elevated levees
along rivers (Clapperton, 1993: 196; Han-
agarth, 1993). Stenocercus prionotus or S.
Gadus seem unlikely to occur in this area
except possibly in these gallery forests, al-
though Fugler (1986) reported S. priono-
tus (Be 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 distabutone may not over-
lap. Stenocercus prionotus is associated
'S 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.
scapularis (see Distribution Patterns in Stenocercus
prionotus).
'4 See the Appendix, UTA 38048. Michael Harvey
(personal communication) recently obtained speci-
mens of Stenocercus caducus at El] Refugio, a lowland
locality at the southern end of the Serranfa de Huan-
chaca (14°44'S, 61°O1’W).
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
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 dry forests, palm sa-
vannas, gallery 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 caducus is known
from Parque Nacional Noel Kempff Mer-
cado and vicinity in Bolivia (see footnote
14; Harvey, 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). Harvey
(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 (Harvey, 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 dry forests sur-
rounding the Gran Chaco” (Harvey, 1997:
35). The montane wet forests (yungas) of
this area are restricted to ridges high
enough for cloud formation during much
of ie year (generally >1,500 m elevation),
and they are surrounded by deciduous dry
211
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 (Holst, 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 ‘aaclen stood. 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 Kempft 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 very similar to
S. caducus (P. FE. 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.
bo
bo
1
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 peculiar (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 he upper
Amazon, the Rio Marafion. Neither S. acu-
leatus nor any other species of Stenocercus
was obtained during herpetofaunal 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 Gregory Vigle). The absence of
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
fimbriatus and S. prionotus has concomi-
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 very deep postfemoral pockets (Type
5) in both sexes, whereas the postfemoral
pockets are more weakly developed in
specimens from Ecuador (Type 2) Oo) lM
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 these 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. fimbriatus and S. prionotus with S.
aculeatus. Taxonomic recognition of S.
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 Coneideres! herein (e.g., see the:
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle
synonymy of Stenocercus prionotus ), | 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 (“Ophryoes-
soides group” as used herein). I have also
included the three other currently recog-
nized species having these characteristics,
S. erythrogaster (Hallowell), S. dumerilii
(Steindachner), and S. tricristatus (Du-
méril), although these are not known from
Peru or Bolivia and are unlikely to occur
there. Character and distributional data in
the key for S. dumerilii 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, butel am un=
aware of other undescribed species of the
“Ophryoessoides 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 will permit identification of all Pe-
ruvian and Bolivian taxa previously con-
fused with S. aculeatus (e.g., Dixon and
Soini, 1975, 1986 [S. fimbriatus]; Fugler,
1983, 1986, 1989 [S. prionotus]) and S. ir-
idescens (e.g., S. huancabambae 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. aculeatus, S. iridescens, and S. lim-
itaris are definitely known from Ecuador.
213
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 dimorphism 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. Summary geographic distribu-
tions are given for each species as a rough
guide to known occurrences. However,
these should be used cautiously as ancil-
lary information in identifying specimens
because distributions of species are some-
times poorly circumscribed. For greatest
utility the key should be used in conjunc-
tion with illustrations herein and in Cadle
(1991, 1998) and Avila-Pires (1995).
2
a
to
icy)
14
Canthal and supraciliaries forming a pro-
nounced crest that ends in an enlarged,
erect, postsupraciliary that may be dis-
tinctly pointed or blunt. Posthumeral and
postfemoral mite pockets absent (Type 1
rial loxovclay Tua uANNKOS)) 2
Canthals and supraciliaries not forming a
pronounced crest; no enlarged, erect post-
supraciliary. Posthumeral pocket absent
(Type 1) to deep (Type 4). Postfemoral
pocket absent (Type 1) to deep (Type 5)
Enlarged postsupraciliary distinctly pointed.
Two enlarged scales above ear opening.
Tibia approximately equal to thigh length
pan Stee oes Ne Stenocercus dumerilii (Steindachner)
(northeastern Para, Brazil)
Enlarged postsupraciliary blunt. No enlarged
scales above ear opening; tibia distinctly
SIorausre tdaenel (Hove oe
EA ENS | Stenocercus tricristatus (Duméril)
(known only from the holotype, probably
from the state of Minas Gerais, Brazil)
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) __...--.--------
Bet ee RLS tel s Stenocercus scapularis (Boulenger)
(intermediate elevations on the Andean
slopes of central and southern Peru; known
elevations greater than 1,000 m)
Superciliary scales not projecting laterally,
the anterior ones elongate, strongly over-
lapping. Midbody dorsal scale rows fewer
than 55 (30-53). Postfemoral pocket vari-
ANS (kyae Il; 2, O; Cir &) 2 4
Posthumeral mite pocket deep (Type 4) and
with an associated scaly flap extending
from its anteroventral border and partally
COMCE AMIN GPity sete ate see ee 5
Posthumeral mite pocket absent to deep
(Type 1, 2, 3, or 4) but without an asso-
Clatealet apy piesa lean eien mas MET eee A hol 5S 6
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
'S 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 cor ated with body size
Aaa thus develop with age. The number of dorsal
scale rows and the development of the postfemoral
pocket are useful clues for subadults.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
scales om each side
ee ee Stenocercus prionotus 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-
largedsand monprojecting seen
myftiea See FS Ree ON 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.'® Seales
between vertebral and dorsolateral crests
smooth or occasionally very weakly keeled.
Let Dee Dds BS Stenocercus fimbriatus 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. 420. 1.0 0 Mie seit et tad vere i
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 wit! ano As, Wee ehtded deel ereecermet 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.
Bb neta eatin. ee 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 Realeer on the shank are rela-
tively more prominent in juveniles than adults.
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA * Cadle
porals and the posterior dorsal head scales.
Two subequal canthal scales on each side.
Head scales keeled, at least posteriorly 9
Two projecting bladelike angulate temporals
in line with superciliary row. A single can-
thal 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
(co) Gaveredhy Ievslkevl ee ee
peste) conte Stenocercus erythrogaster (Hallowell)
(northern Colombia)
Posthumeral and postfemoral pockets deep
in both sexes (Types 4 and 5, respective-
ly).!7 Anterior gular scales smooth to weak-
IWakeele diye iw te wees Sol eh Pat ee ee 10
10. Interparietal 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)
Interparietal 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. ___.
PEER athe Stenocercus limitaris 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).
ae Tu 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
GRES|O) (CLIN GONE Bo cee
densa NRTA Stenocercus huancabambae Cadle
'7 An undescribed species in the next 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 ancillary data for identification.
215
(dry inter-Andean valleys of the upper Rio
Marafion in Cajamarca and west central
Amazonas departments, northern Peru)
ACKNOWLEDGMENTS
Loans and other assistance were facili-
tated by Linda Ford, Darrel 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 P. 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.
huancabambae and S. limitaris) 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 Métraux’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 Heller’s collec-
tion; Charles W. Myers (AMNH) did the
same for Harvey Bassler’s journeys in
northern Peru and provided the base map
used to prepare Figure 1; Robert S. Voss
216
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
(S. huancabambae, 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
[ea. 1,000 m; 01°59’S, 77°58’W] (USNM 200882-84).
[PProv. Napo/TuncurAnuA]: Llanganates area!’
(FMNH 23527). Prov. Pastaza: Rio Pastaza, Abita-
gua [01°23'S, 78°05'W] (FMNH 25803-05, 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 Bobonaza [314 m; 02°04’S,
76°58'W] (USNM 200892). [PPRov. PASTAZA]: Banos,
Mera Trail [? = between Banos and Mera"; approx-
imately 01°30'S, 78°10'W] (FMNH 28012).
PERU: [Depto. LA LipertaD]: E Peru, Pampa
Seca, Rio Mixiolla [=Rio Mishollo] Valley, Upper
Huallaga region, 4300 ft [2, 1,311 m; approximately
08°16'S, 76°58'W]?) (AMNH 57085). [DEPpro. Lor-
18 T 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, 1993). The locality may refer to lower ele-
vations in this range.
'° Bafios is a famous collecting locality at the foot
of Volcan Tungurahua 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 Bajos and Mera, which
is at 1,160 m. See Brown (1941) and Chapman (1926)
for discussion.
°° Harvey Bassler collected Stenocercus for the
AMNH at two localities on the Rio Mixiolla (=Rio
Mishollo): Pampa Seca and La Pinita (see Stenocer-
cus fumbriatus), 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 |
Piflita, respectively, in extreme southeastern La Lib-
ertad Department, as indexed by Stiglich (1922). —
Both localities are in Ongén District and the coor-
dinates given are those for the town of Ongon. Stig-
lich (1922) states that Pifiita is a small village on the ©
Quebrada Pedernal, a left tributary of the Rio Mish-
ollo. Apart from indicating that Pampaseca is a farm, |
NEW SPECIES OF STENOCERCUS FROM PERU AND BOLIVIA ° Cadle
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 “Icu-
to Cuesta” by Lamas, 1976] (AMNH 56413).
Stenocercus caducus
BOLIVIA: No SPECIFIC LOCALITY: _BMNH
1946.8.29.76 (holotype of Leiocephalus 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). DErro. 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:2! 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). [| Provincia |
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, Rfo 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, 61°15’W] (UTA
38048). [DEepro. Tara]: Villa Montes [21°15'S,
63°30'W] (MCZ 28634). Misi6n San Francisco
[21°15’S, 63°30'W] (BMNH 98.7.7.5; specimen col-
lected by Alfredo Borelli, whose San Francisco = Vil-
la Montes fide Paynter, 1992).
PARAGUAY: [DEpTO. CAAaGuAzv]: Pastoreo [ap-
proximately 25°23'S, 55°52’W] (MCZ 34214—-15).
[DEPro. CENTRAL]: Asuncién [25°16’S, 57°40'W]
(FMNH 9496). Colonia Nueva Italia [25°37’S,
57°30’W] (FMNH 42281).
Stenocercus erythrogaster
COLOMBIA: [DEptTo. MAGDALENA]: Rio Frio
[30-450 m; 10°55'N, 74°10’W] (MCZ 29707). Santa
Marta 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.
*1 The specimen was collected in November or De-
cember 1913 by José Steinbach, who collected at a
locality known as “Las Juntas” during that same pe-
riod (Paynter, 1992). The two localities are assumed
to be the same. The variant spelling “Yuntas” does
not appear in any sources consulted.
bo
7
Stenocercus fimbriatus
PERU: No SPECIFIC LOCALITY: (FMNH 56070).
DeEpro. HUANUCO: ca. 35 km NE Tingo Maria, Heda.
Santa Elena, ca. 1000 m [approximately 08°57’S,
76°02'W] (LSUMNS 26966-67). Approximately %
mile E Universidad Agraria de La Selva, Tingo Maria,
vicinity of Rio Huallaga [3, 09°18’S, 75°59'W],
USNM 193684. [DEpro. LA LIBERTAD]: E Peru, La
Pinita, Rio Mixiolla, tributary 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 Biolégica 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-
DRE DE Dios: Pakitza Station [Rio Manu], Manu Na-
tional Park [4, 11°56’S, 71°17’W] (Victor R. Morales
18235). Drpro. UcayYALi: Rio Curanja, Balta, ap-
proximately 300 m [approximately 10°08'S, 71°13'W]
(LSUMNS 17519, 25402-04, 26720-23). Alto [Rio]
Purtis, 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 scapularis**
PERU: No SPECIFIC LOCALITY: (FMNH 56444).
[DEpPTo. 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-
tin [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.
23 The occurrence of Stenocercus scapula ris 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
(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
bank of the Rio Inambari, 1,830 m, 13°30'S.
70°00'W |? (AMNH 1701).
4 According to the field catalogue in the FMNH
Mammal Division 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 ‘Inambary River.
The country is very broken, with deep narrow can-
ons, and is cover ed with a dense undergrowth of
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 Juliaca ... the Inca Mines are situated
about 200 miles northeast of Juliaca, on the east
side of the Andes, on the Inambary River, a trib-
utary of the Amazon, and at a much lower altitude
than Juliaca. The altitude and geographical position
were correctly given in the fannie 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 the 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Stenocercus sp.
PERU: Deprro. AMAZONAS: 17 km ENE Balsas
[06°49'S, 78°00'W] (ROM 16458).
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ON THE SUBFAMILY XYLOPHAGAINAE
FAMILY PHOLADIDAE, BivatviA, MoLtusca)
RUTH D. TURNER
HARVARD UNIVERSITY mote ~ VOLUME 157, NUMBER 4
CAMBRIDGE, MASSACHUSETTS, U.S.A. 31 October 2002
(US ISSN 0027-4100) . |
OR DISTRIBUTED BY THE
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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 Emest 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.
es of the New England Zoological Club 1899-1947. (Complete
sets only.)
Proceedings of the Boston Society of Natural History.
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© The President and Fellows of Harvard College 2002.
ON THE SUBFAMILY XYLOPHAGAINAE
(FAMILY PHOLADIDAE, BIVALVIA, MOLLUSCA)
RUTH D. TURNER’
CONTENTS
BicitonallNotes int © ean eee 223
NUNS trac ty. ea RAAT Ra Sea ars Betleel pee Rh er 223
TO LUG LOI ie een eaves Weare ee ae ete 224
Species Groups in the Genus Xylophaga ___. 225
\V GREET OS ypc is ei, a a ae 227
Preliminary Report on the Results of
Experiments on the Ecology of Deep-
Sea Wood Borers and the Role of Wood
Hhay ove} IDYStS50) (SSE Lene eee ee 234
SOUT ey, ecb ee ee 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 Taki and Habe _. 241
Xylophaga whoi Turner new species 242
Xylophaga profunda Turner new species — 243
Xylophaga abyssorum Dall 245
Xylophaga duplicata Knudsen __- 247
Xylophaga muraokai Turner new species — 247
Xylophaga atlantica Richards 249
Xylophaga washingtona Bartsch 250
Xylophaga rikuzenica Taki and Habe _... 252
Xylophaga depalmai Turner new species 253
Xylophaga guineensis Knudsen 256
Xylophaga mexicana Dall __..........----.---------- 257
Xylophaga tipperi Turner new species __. 259
Xylophaga bayeri Turner new species __.. 260
Xylophaga japonica Taki and Habe 261
Genus Xylopholas Turner 1972 _............-...-.- 262
Xylopholas altenai Turner 263
Genus Xyloredo Turner 1972 264
MPIDRBOID) TOGA VOTRE 265
Xyloredo ingolfia Turmer —_.....------------------ 266
Kyloredownaceli Tamer 2-22) 267
Achenexalercles tea ee ee ee 268
Witeratuner© ite cern een res arene ner mee 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’, 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
with the intent of readying it for formal publication. Two
outside authorities, Dr. Jorgen Knudsen of the Zool-
ogisk Museum, Kgbenhavns Universitet, Kobenhavn,
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 incorporated 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 Dore)
bo
bo
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-
nificantly 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 Xylophaga 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, S40 Tomé, Gulf of Guinea.
However, the Sio Tomé 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.
4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Until recently, the Xylophagainae were
considered to be deep-sea organisms of lit-
tle 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-
terbury 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- |
dinidlike burrow that is lined with a cal-_
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 Xylophaga dorsalis Turton, and
Turner (1955) for X. atlantica.
SPECIES GROUPS IN THE
GENUS XYLOPHAGA
Genus Xylophaga 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
XYLOPHAGAINAE ° Turner 925
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:
bo
Text-Figure 1.
siphonal retractor
pedal retractor
posterior adductor
ventral adductor
umbonal-ventral ridge
Bulletin Museum of Comparative Zoology, Vol. 157,
umbonal reflection
extended.
Group 1.
Group 2.
Mesoplax composed of two
simple flat or slightly curved 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.
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.
wolfft Knudsen, X. hadalis Knudsen, X.
galatheae Knudsen, X. murrayi Knud-
sen, X. africana Knudsen, X. panamen-
sis Knudsen, and X. clenchi Turner and
Culliney.
Group 3.
Group 4.
No. 4
anterior adductor
incurrent
siphon
umbonal- ventral sulcus
mantle
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
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
usually with small cirri on both open-—
ings. Group 3 includes X. supplicata |
Taki and Habe, X. lobata Knudsen, X.
tubulata Knudsen, X. bruuni Knudsen, |
X. obtusata Knudsen, X. whoi Turner n. |
sp., and X. profunda Turner n. sp. |
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.
Mesoplax composed of two.
}
Group 5. Mesoplax composed of two
plates that are more or less triangular in
outline, with a ventral portion ranging
from 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. depalmai Turner n. sp.,
X. guineensis Knudsen, X. mexicana
Dall, X. tipperi Turner n. sp., X. bayeri
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 of Xylophaga
XYLOPHAGAINAE * Turner ORT
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 Xylophaga 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 material 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 from 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-
iN)
bo
8 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
dorsalis
a praestans
0 A
0 grevei
bruuni
indica Q
tomlini or
o
erecta
Rm:
rikuzenica
supplicata
japonica
tubulata
clenchi
atlantica
° abyssorum
depalmai
washingtona S
pir aoka proto
. Sue
obtusata
duplicata .
guineensis
africana
Oe
wolffi
ct ~obata —___
murrayi ae o' wliata
concava
aurita
turnerae
AS
galathea
uy 27
hadalis
globosa
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 exhibits variation that possibly
is genetic because the substrate and all
other parameters were as nearly uniform
as possible, and the third case illustrates
variation in a normal growth series.
Variation Due to Different Substrates.
Variation in Xylophaga washingtona in re-
sponse to the substrate can be denounce at=
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).
mhes2)xe6r< 0) d-inl (0S P52 alo a7—
mm) 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
which the borer larvae settled and into”
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.
Ihe 1g 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-
beau 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
specimens had not greatly increased in di-_
XYLOPHAGAINAE * Turner 229
TABLE 1. VARIATION OF XYLOPHAGA WASHINGTONA BURROWS IN DIFFERENT TYPES OF WOOD.
No. Burrow Burrow
Specimens Length Diameter
Wood eanined (m m) (mm) Remarks
Cedar 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.
Ash 125 21.0 5.0 many dead, burrows running together, panel
heavily attacked, particularly at one end.
Maple 75 11.0 5.5 well distributed, with a little more concentration
around the hole at one end.
Pine 35 15.0 5.0 clustered at one end, newly settled to adult.
Oak +50 15.0 6.0 evenly distributed, shells yellowish-green from
wood.
Fir 46 12.0 4.5 many newly settled, often cut into burrows of oth-
er specimens.
Redwood 46 14.0 4.5 specimens stained dark red brown by wood.
Greenheart 75 2.75 1.75 concentrated around edges.
Afambeau 19 0.75 0.05 concentrated at one end, all very small, many in
umbo stage veliger, 15 small depressions with-
out animals.
Antidesma pulvinatum 150 0.10 0.05 many newly settled, just beginning metamorpho-
sis.
= [+__—____|
1mm
1mm
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
Maple
Oak
5mm
Fir
Text-Figure 5. Typical specimens of Xylophaga washingtona collected from submerged oak (top) and fir (bottom).
XYLOPHAGAINAE *° Turner 231
Redwood
Greenheart
Afambeau
ba Saar
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
eae Sa Sy
0.1mm
Text-Figure 8. Typical specimens of Xylophaga washingtona collected from a submerged panel of Antidesma pulvinatum.
ameter as they bored so that the larval
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-mm
scale bar, and are arranged in order of de-
creasing size. The number of denticulated
ridges on the anterior slope of these spec-
imens varies from 12 to 20. Specimens
from redwood, greenheart, Afambeau, and
Antidesma pulvinatum (Text-Figs. 6-8)
were drawn to their own scales and, al-
though these specimens are much smaller,
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 muscle in response to in-
creased activity of boring. The general
shape and sculpturing of the muscle 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 material 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
NL
Boe
Phenolic Laminated Rod
XYLOPHAGAINAE ° Turner 233
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 removed 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 of 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
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. bayeri 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.
bayeri (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 ee
eral arms produce an elaborately lobed
mesoplax. Nothing is known of the allo-
metric changes ihe take place during
growth of ine peculiar dorsal plates of xX
tubulata 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 laboratory. In the meantime,
it seems best to consider these forms as
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
members of growth series rather than dif-_
ferent species, particularly because the se-_
ries in each case was built up from mate- i
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)—Alvin 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-
morphosing X ylophaga with one to two
rows of denticulated ridges were removed
from the panel. The specimens were ap-
proximately 300 ym 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 tose 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
t
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
lar: ge 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 with 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 measured § ae in length; others
measured 40, 33, 32, 30, and 24 mm. It is
obvious that the se were finding suffi-
cient food either in or on the wood B grow
at a fairly rapid rate.
The first young crabs to find the wood
may have fed on the newly settled Xylo-
phaga larvae and this might explain the
patchy distribution of tne 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 length to get
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
XYLOPHAGAINAE ° Turner 935
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 carrying larvae on the umbonal area
of the valves. The larvae measured 0.2 mm
in length. The large species of Xylophaga,
X. profunda Turner n. sp., had not been
seen before. The valves were 14 mm in
length, and one specimen measured 40
mim 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
che 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 X ylophaga
burrows and on the surface of the wood,
I also found two other polychaete worms.
One belonged to the family Chrysopetali-
dae and the other to a family of polynoid
worms.
A preliminary 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
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) Xylophaga 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 larvae of X. profunda n. sp.
were settling in early March. 3) Xylophaga
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 Xylophaga 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 lenovell
edge, this isthe 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
Xylophaga concava Knudsen
Plate 1
Xylophaga 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; paratype, 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
umbos. Siphons nearly the same 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 Specimen:
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, Colom-
bia, whereas station 739 is about 90 miles.
west of Ensenada Guayabo, Panama. A
single specimen of X. concava was taken’
by the R/V Pillsbury at station 526. This
locality is about midway between the two
Galathea stations. |
The Pillsbury 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 them 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.
Xylophaga concava is closely related to|
Xylophaga g gerda Turner n. sp. but differs
in the size Ok the mesoplax, the type of |
siphonal openings, and the chimney. (See
Remarks under x. gerda.) Xylophaga con-.
vava also is related to X. erecta Knudsen:
(1961) from the Sulu Sea. Knudsen (1961) |
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)
specimens and unfortunately no further
records have been obtained.
Range. From off Ensenada Guayabo,
Pannen south to off the Gulf of Tibuga,
Colombia, in depths from about 915 “he
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); Pillsbury, 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 ee
(5°49'N, 78°52’W) in 3,270-3,670 m.
Xylophaga gerda? Turner new species
Plates 2, 3
Holotype. 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
curved 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
Welate 2, Fig. 7).
Description. Shell globose, fragile,
reaching 3.0 mm in length and 2.8 mm in
height, alae 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-
2 Named for R/V Gerda, Rosenstiel School of Ma-
rine and Atmospheric Sciences, University of Miami,
Miami, Florida, whose station 499 is the type locality.
XYLOPHAGAINAE * Turner 237
TABLE 2. MEASUREMENTS OF XYLOPHAGA GERDA.
Length Height
(mm) (mm) Location
Tail 1.0 Gerda, station 266
15 1.4 Gerda, station 266
1.8 2.0 Gerda, station 266
2.5 2.3 paratype
3.0 2.5 holotype
3.0 2.8 Pillsbury, station 328
3.3 3.0 Pillsbury, station 944
3.8 3 Pillsbury, station 944
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 brown periostracal cylin-
der containing fecal material may extend
nearly one hale the length of the excurrent
siphon (Plate 2, Figs. 124) Chimney built
in sections, composed of fine fecal material
agelutinized 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 very 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
(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: Pillsbury, station 944, 45 miles N of
Port Louis, Guadeloupe Island (16°32.2'N,
61°36.8'W) in 364-421 m. PANAMA: Pillsbury, sta-
tion 328, about 25 miles N of Punta San Blas, Gulf
of San Blas (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 Ceram [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 me-
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-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
TABLE 3. MEASUREMENTS OF XYLOPHAGA GREVEI. |
Length Height
(mm) (mm) Location
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 umbonal 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
specimens of X. grevei. They are much)
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)
only two specimens, also was from Gala-
thea, station 444. It has valves, muscle |
sears, 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-
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 clenchi? Turner and Culliney
Plates 5-8
Xylophaga clenchi 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.
XYLOPHAGAINAE ° Turner 239
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 where 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
TABLE 4. MEASUREMENTS OF XYLOPHAGA CLENCHI.
Length Height
(mm) (mm) Location
Des 2a Ingolf, station 67
B.D 2.0 Atlantis II, station 124
3 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 Pillsbury, 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 fragmentary 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. wolfft 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. swpplicata Taki and Habe.)
Xylophaga clenchi Turner and Culliney
also appears to be closely related to X.
panamensis Knudsen but the latter differs
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 |
in having the plates of the mesoplax |
smooth. 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. panamensis are unknown. |
The variation in the mesoplax 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 which |
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 R/V 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 |
from 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 R/V |
Atlantis IT 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 very delicate. This raises the_
question as to whether or not at least some
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 larvae, 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 m; off Cape Hatteras (35°44'N,
75°15'W) in 35 m. GEORGIA: Pillsbury, station 104,
about 80 miles SE of Brunswick (31°00'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 the 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
XYLOPHAGAINAE ° Turner 241
TABLE 5. MEASUREMENTS OF
XYLOPHAGA SUPPLICATA.
Height
(mm)
8.5 8.8
Length
(mm) Location
paratype MCZ 194820
4,862 m:; Atlantis II, station 119, just S of Bermuda
Pillsbury, station 328, about 25 miles off Punta San
Blas, Gulfo San Blas (9°55.8'N, 78°59.8’W) in 420—
640 m. VENEZUELA: Pillsbury, 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
Metaxylophaga supplicata Taki and Habe, 1950, II-
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
mesoplax.
Xylophaga bruuni 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 necessary to
obtain more material before a definite
statement can be made concerning the sta-
242
tus of X. bruuni. (See also Remarks under
Xylophaga whoi Turner n. sp.)
Specimens Examined. JAPAN: Tosa Bay, Shikoku,
in 100 fathoms (183 m).
Xylophaga who/* Turner new species
Plates 10, 11
Holotype. 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 sharp 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, Shan
fragile, and with a very thin, light tan per-
iostracum. Pedal gape angle walvoue 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 fhe
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
‘An acronym for Woods Hole Oceanographic In-
stitution, whose research vessel Atlantis collected the
holotype.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
TABLE 6. MEASUREMENTS OF XYLOPHAGA WHOI.
Length Height
(mm) (mm) Location
2.5 ORS) Gerda, station 266
3.1 3.0 Gerda, station 266
6.9 6.2 holotype
7.0 6.8 Pillsbury, station 394
Me 7.0 Pillsbury, station 944
9.5 9.0 Pillsbury, station 944
12.5 11.5 Pillsbury, station 944
13.8 13.1 Pillsbury, station 944
14.9 14.0 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 heen 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 thes excurrent
siphon small ana with a few fine cirri on
each side ventrally.
Measurements. 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. tub-
ulata Knudsen from Macassar Strait, X.
bruuni 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-
ma). Xylophaga whoi Turner n. sp. is the
first tubulate species reported from the At-
lantic. It is most closely related to X. bru-
uni Knudsen from the Mindanao Sea but
differs in that the mesoplax has a propor-
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
the shell but increases rapidly to a large
condyle. Xylophaga 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 from 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. bruuni 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 R/V Pillsbury 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. bruuni. 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
XYLOPHAGAINAE ° Turner 943
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 larvae 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. twbulata 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, FLOR-
IDA: Gerda, station 266, off Fowey 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: Pillsbury, station 944, 45 miles
N of Port Louis, Guadeloupe Island (16°32.2'N,
61°36.8S'W) in 364-421 m. COLOMBIA: Pillsbury,
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
Holotype. 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 segmented.
Description. Shell globose, reaching
about 11 mm in length, thin, fragile, and
QA4
TABLE 7. MEASUREMENTS OF
XYLOPHAGA PROFUNDA.
Length Height
(mm) (mm) Location
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
95 88 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 reflec-
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 giving
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. abyssorum Dall but differs in
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
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 mesoplax 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. lobata
Knudsen from the Sulu Sea. Xylophaga
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 intervening 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 re 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. whoi
and X. supplicata, both of which have a
single large tooth. Unfortunately, Knudsen
digh 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,
probably of the same length and capable
of complete retraction ation 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 suum cee in the Tongue of the
Ocean at 25°54'N, 77°49’'W, Aa 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 m; Tongue of the Ocean,
Tower 3 (24°53. 2'N, 77°40. 2'W) in 2 5066 m.
Xylophaga abyssorum Dall
Plates 14-16
Xylophaga abyssorum Dall, 1886, Bulletin Museum
of Comparative Zoology, 12: 317, pl. 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, pl.
93.
Distinctive Characters. Valves with a
prominent 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
XYLOPHAGAINAE ° Turner IA5
toward the ventral margin in older speci-
mens. Umbonalventral 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 ateached by a
periostracal fold. lWnibes inflated, particu-
larly in young specimens.
Inner surface of valves white and glazed.
Muscle scars well marked. Posterion 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 variables with a well-devel-
oped, more or less triangular basal portion,
occasionally with lateral notches in young
specimens. Dorsal portion developing and
varying with age; in young specimens, con-
sisting of lobes extending anteriorly from
the posterior ends of they plates (Plate 14,
Fig. 2) followed by lobed lateral folds
(Plate 14, Figs. 3-5, 7), which grow and
eventually eodlesce 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
TABLE 8. MEASUREMENTS OF
XYLOPHAGA ABYSSORUM.
Length Height
(mm) (mm) Location
3.0 2.6 Pillsbury, station 944
3.0 2.9 Tongue of the Ocean, off NE tip of
Andros Island, Bahama Islands
Pillsbury, station 944
Pillsbury, station 944
holotype
Gerda, station 266
Pillsbury, station 944
3.3 3.0
Se) ei
4.0 3.5
4.0 3.6
5.4 4.5
Measurements. See Table 8.
Remarks. 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. abyssorum 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;
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
the average measurement of the young |
was 0.3 mm in length.
Xylophaga abyssorum 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
muscle scars of X. abyssorum are well
marked. In the young stage, the mesoplax
of X. abyssorum is similar to that of X. bay-
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
GO soi,
}
I)
|
|
|
f
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: Pillsbury, station
944, 4.5 miles N of Port Louis, Guadeloupe Island
(16°32.2'N, 61°36.8'’W) in 360-420 m.
TABLE 9. MEASUREMENTS OF
XYLOPHAGA DUPLICATA.
Length
(mm)
Height
(mm) Location
Bia 3.5 holotype
2) 5.0 Gulf of Tehuantepec, Mexico
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 sculptured 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 Gulf 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
XYLOPHAGAINAE ° Turner IAT
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 1O9 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 I-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 unequal length, the ex-
current slightly shorter and with 8-10
prominent 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 sculptured 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
depressed and sculptured with fine growth
lines. Disc and posterior slope sculptured
with growth lines only.
Inner surface of valves smooth and shin-
ing. Umbonal-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 ductor muscle scar
long, narrow, and traversing the ventral
endl of the bonalevenerall 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, curved, wedge-shaped
shield, fitting closely against the surface of
the valves. Posteriorly, the mesoplax curves
upward and forward 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
TABLE 10. MEASUREMENTS OF
XYLOPHAGA MURAOKAI.
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
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 thokened 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.
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 AR ennettor 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 sculpture 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
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-
XYLOPHAGAINAE *° Turner YAY
TABLE 11. COMPARISON OF THE U.S. NAVAL CIVIL
ENGINEERING LABORATORY TEST SITES (AVERAGE
VALUES TAKEN FROM MURAOKA, 1967).
Tempera- Dissolved Hydrostatic
Depth ture Oxygen Pressure
Locality (ft) (EG) (ml/L) (psi)
Test Site I 5,640 2.5 1.26 2,482
Test Site IT 2,370 5.0 30 1,043
lands); STU I-1 at 5,300 ft (1615.4 m) exposed March
1962—February 1965; STU I-2 at 5,640 ft (1,720 m)
exposed October 1963-—October 1965; STU I-4 at
6,800 ft (2,072.6 m) exposed June 1964—7 July 1965;
STU I-5 at 6,000 ft (1,828.8 m) exposed 25 August
1965-12 February 1966.
Xylophaga atlantica Richards
Plate 20
Xylophaga atlantica Richards, 1942, Nautilus, 56: 68,
pl. 6, fig. 4 (east coast of the United States). Ho-
lotype, Academy of Natural Sciences Philadelphia
178741; Turner 1955, Johnsonia, 3(34): 152-154,
pl. 91, figs. 1-6 (type locality, Mount Desert Island,
Maine). (See Turner, 1955: 153.) [Not Turner,
1954, Johnsonia, 3(33): 5-6, pl. 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 margin
and meet at a broadly obtuse angle. Ven-
tral portion 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. Excurrent siphon
slightly shorter than the incurrent, the ex-
current aperture with 15—20 large papillae,
incurrent aperture with a double row of
numerous minute papillae (Plate 20, Fig.
83),
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 bottom one
had 85 specimens of X. atlantica, the one
250
at 150 ft had six specimens, and the one
at 100 ft had only one specimen. This sug-
gests that, like X. washingtona Bartsch, X
Feet Turner n. sp., x depalmai Tain:
er n. sp., and probably all Xylophaga, 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 Thewe 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
spac stouter papillae (Plate 20, Figs. 1,
Xylophaga atlantica is oviparous and
does not brood its young. The reproduc-
tion and larval development of X. atlantica
are fully discussed by Culliney and Turner
(1976); they detail methods of laboratory
culture and illustrate the various larval and
growth stages. A reexamination of the
specimen reported by Turner (1954: 5-6,
pl. 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
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 ndameton |
scar is irregularly marked, whereas that of |
X. washingtona has regular herringbone
markings.
Range. From Newfoundland south to
Cape Henry, Virginia, in depths ranging
from about 15 to 1,242 m.
Specimens Examined (New Records Since Turner,
1955). CANADA, NEWFOUNDLAND: % 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 pa anels). 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, pl. 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, 64-115.
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.
Locality Depth (m) Exposure Reference
3 miles off Oceanside, California 109 4 months Turner, 1956
STU* II (1 and 2), California 722
STU* I (1-4), California 1,615—2,066
19 miles off Depoe Bay, Oregon 200
25 miles off Depoe Bay, Oregon 500
40 miles off Depoe Bay, Oregon 1,000
* STU, submersible test unit.
bone markings. Proximal end of the com-
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 having 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
fibe extending between the lateral ridges,
and apparently lacking papillae (Plate. OL.
igs.) 125)'3); Ineur tent 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. washingtona, ane inter-
esting observations 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 larval and
adult X. washingtona 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
6.5, 13.4 months
4, 13, 25, and 35
months
38 and 72 days
2 months
2 months
Muraoka, 1965, 1967
Muraoka, 1964, 1966a,
1966b, 1966c
Tipper, 1968
Tipper, 1968
Tipper, 1968
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, % mile west of Hermosa
Beach, Santa Monica Bay, California. This
is the shallowest record known for the set-
tlement of larvae. 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: Breen
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 Moca 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 aethinrent
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 larvae of X.
washingtona off Depoe Bay, Oregon, was
dence 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
bo
bo
te)
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 1O—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,
IS mm) in fir, and 0:5 mim 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 observations 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 I-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 ie 1 exposed fork 35 months had spec-
imens with tunnels reaching 35 mm in
length. These tunnels averaged 10 mm in
ghoumeter 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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
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
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°59’ 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, % 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 L.,
Santa Barbara Islands (34°06’N, 120°42’W) in 722 m;_
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, foe % mile E of Empire Landing, Santa
Catalina Island, Santa Barbara Islands (33°25' 50"N, |
118°24'50"W) in 84 m (Allan Hancock Foundacon
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 Adductel
muscle scar elongate oval, tapering dorsal-|
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) Plated 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. aurita 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 hay-
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.
XYLOPHAGAINAE ° Turner 253
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 &§ 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 curved dorsal margin
that is not reflected.
Inner surface of valves smooth and glis-
tening. Umbonal—ventral ridge narrow,
high, and not enlarging at the vente: al mar-
gin to form a able 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 John DePalma, U.S. Naval Oceano-
graphic Office, who was responsible for the tests off
Fort Lauderdale, Florida; in the Tongue of the
Ocean, Bahama Islands; and in Penobscot Bay,
Maine.
254
TABLE 13. MEASUREMENTS OF
XYLOPHAGA DEPALMAI.
Length Height
(Gaaaa (mm) Location
6.0 5.5 off Fort Lauderdale, Florida
7.6 7.0 off Fort Lauderdale, Florida
8.0 WED 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,
irregularly oval, located posterior to the
umbonal—ventr: all 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 suintace
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 Re 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 surface 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 Bre: irregular, glasslike granules
(Plate 23, Figs. 1, Ay 5),
IMeaerements See Table 13.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 |
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 hay-
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,
ich it also somewhat resembles, X. dem
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
ines siphons. |
Variation exhibited in the mesoplax of X.
depalmai is unusual and unexpected, for.
in most species of Xylophaga the mesoplax,
is remarkably uniform unless the specimen:
is injured or stenomorphic. 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 Turner n. sp. was)
the most common species occuring 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
XYLOPHAGAINAE ° 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 Depth of
Shore Depth Panel
Panel no. (miles) in Feet (m) in Feet (m)
2 panels without 2.5 300 on bottom
numbers (91.44)
Ww44 1.8 100 on bottom
(30.48)
H50 2.3 200 100
(60.96) (30.48)
H56 23, 200 on bottom
(60.96)
G62 2.6 300 2.00
(91.44) (60.96)
G68 2.6 300 on bottom
(91.44)
59 2.3 200 on bottom
(60.96)
492, 3.2 500 2.00
(152.4) 60.96)
493 By 500 200
152.4) 60.96)
497 3.2 500 300
152.4) 91.44)
498 Bi) 500 300
152.4) 91.44)
500 3.2 500 400
152.4) (121.92)
D301 3.2 500 400
152.4) (121.92)
502 BP 500 400
(152.4) (121.92)
504 3.2 500 on bottom
(152.4)
505 BP) 500 495
(152.4)
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, more wood was
available on the bottom in this vicinity,
supporting a native population and a ready
source of larvae. However, this also may
nae fa eee
Sept. 1961 June 1962 +250
Jan. 1964 Oct. 1964 1
Jan. 1964 Oct. 1964 2
Jan. 1964 Sept. 1964 5
Jan. 1964 Sept. 1964 6
Jan. 1964 Sept. 1964 31
24 Jan. 1964 4 Jan. 1965 3
20 Oct. 1965 13 Oct. 1965 3
20 Oct. 1965 13 Oct. 1966 10
20 Oct. 1965 13 Oct. 1966 5
20 Oct. 1965 13 Oct. 1966 il
20 Oct. 1965 13 Oct. 1966 3
20 Oct. 1965 13 Oct. 1966 2
20 Oct. 1965 13 Oct. 1966 3
20 Oct. 1965 13 Oct. 1966 8
20 Oct. 1965 13 Oct. 1966 6
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°
Ghat 100i mand) frome(itos95~ Cratel ds
m. 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
miles W of Tampa, OTEC station 3089 in 126 m; 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-
gle’s 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
Xylophaga 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 2. mm long. Uinbonalvene
tral sulcus only slightly impressed, bound-
ed anteriorly by a low, inconspicuous,
rounded ridge and posteriorly by a narrow,
well- deaned ridge, occasionally almost
bladelike. Disc and _ posterior slope
smooth.
Inner surface of valves smooth and glis-
tening. Muscle scars barely visible. Poetes
rior Ralductor 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
TABLE 15. MEASUREMENTS OF
XYLOPHAGA GUINEENSIS.
Length Height
(mm) (mm) Location
0.9 Pillsbury, station 260
1.3 NED, Pillsbury, station 260
1.3 1.5 paratype (estimated from Knudsen,
1961, fig. 38b)
1.5 1.3 Atlantique Sud, station 33 |
Q® G9) 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! from a
piece of wood; not a single complete shell
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 mesoplax.
and the large sculptured, bright yellow
prodissoconch.
In the course of examining wood |
dredged from off the western boast of Af)
rica, T obtained four dead specimens, one
with a complete mesoplax (Atlantique Sud,
station 33), and seven small living speci-|
mens, 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 symmetrical.
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.
Although stenomorphs are commonly
found in Xylophaga (see section on Vari-
ation), the specimens dissected from the
wood dredged by the Atlantique Sud Ex-
pedition and the Pillsbury 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 mesoplax, the strength of the
ridge bounding the ara bomal-rentiel 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%.
However, the fringe on the lappets could
only be detected under a compound mi-
croscope at 430. 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. Xylophaga 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 extra 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
XYLOPHAGAINAE ° Turner 257
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: Pillsbury,
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°03'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 Dall
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, pl. 90.
Distinctive Characters. Umbonal—ven-
tral sulcus bounded posteriorly by a nar-
row, sharp 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, ane 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 recurved anteriorly, ap-
258
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
near ly 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
flection. Chondrophore and internal liga-
ment well developed.
Mesoplax somewhat variable but usually
slightly longer than wide, with the medial
margin of its 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 eomeition 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
covering the umbonal re- -
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
TABLE 16. MEASUREMENTS OF
XYLOPHAGA MEXICANA.
Length Height
(mm) (mm) Location
4.8 4.5 Malibu Reef, California
5.0 4.2 holotype
UD ES Malibu Reef, California
8.0 TS 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 dubium but this would necessitate de-
scribing the Californian specimens. Be-
cause ihe 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 g alahoce 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 (Dall, 1909), he gave the
range of X. globosa Sowerby as Go 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. | have not seen a specimen of |
globosa from north of Chiloe Island,
Chile. The valves of X. duplicata some- |
what resemble those of X. mexicana; 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.
mexicana 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.
mexicana 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 serve 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
Xylophaga. 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 Turner new species
Plate 29
Holotype. MCZ 316736; paratype, MCZ
Sl6V3S7.
Type Locality. 3.2 miles off Fort Lau-
derdale, Florida (26°04’N, 80°04’W) in
152.4 m (500 ft) ina 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
XYLOPHAGAINAE ° Turner 259
embedded along the side and with knobby
pustules at the posterior end.
Description. Shell globose, reaching 9
mm in length and §.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 smooth, 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 sharp 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
TABLE 17. MEASUREMENTS OF XYLOPHAGA TIPPERI.
Length Height
(mm) (mm) Location
2.0 2.0 off Fort Lauderdale, Florida
4.6 4.] off Fort Lauderdale, Florida
UD 7.0 off Fort Lauderdale, Florida
8.2 Ts) 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. depalmai
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 incurrent siphon of X. tipperi
is papillose. Xylophaga bayeri 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.
6).
Only eight specimens of X. tipperi Turn-
er n. sp. were obtained. They were all
from panel 505 and were collected along
with X. depalmai Turner n. sp. The spec-
imens, even the smallest, were consistent
in having a broad, compressed mesoplax as
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
well as in the characteristics of the si- |
phons. For data on the testing site, see in-
formation under X. depalmai.
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 bayeri’ Turner new species
Plates 30, 31
Holotype. MCZ 316738; paratype, MCZ
316739.
|
}
{
i
|
|
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 from 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, sharp ridge. Posterior slope
rather low and sculptured with well-
marked growth lines.
7 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 Pillsbury and Gerda.
TABLE 18. MEASUREMENTS OF XYLOPHAGA BAYERI.
Length Height
(mm) (mm) Location
8.0 EO holotype
4.2 4.0 off Mona Island, Puerto Rico
5.4 4.9 Gerda, 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 8.2 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
XYLOPHAGAINAE ° Turner 261
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; Gerda,
station 266, off Fowey Rocks (25°39'N, 79°58’W) in
about 340 m. 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
TABLE 19. MEASUREMENTS OF
XYLOPHAGA JAPONICA.
Length Height
(mm) (mm) Location
13.2 10.8 holotype (from Taki and
Habe, 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
ron area, extending over the umbos,
ear-shaped, moder ately inflated, with a
narrowly rounded periphery and distinct
concentric sculpture (Plate 32, Figs. 2-5).
Siphons capable of extending two to three
times the length of the Hol, Excurrent si-
phon about one third the length of the in-
current, lappets extending along the dorsal
surface of the fourm 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 Hoe salis 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
the umbonal-ventral sulcus more pro- |
nounced.
Unfortunately, the material from off |
Java and South Burma noted in the re-
cords below was poorly preserved, 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 Bruun 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,
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 Bruun, station 23, about
77 miles W of Clara Island, South Burma (10°30’N,
96°35'E) in 384 m. :
GENUS XYLOPHOLAS TURNER 1972
Xylopholas Turner, 1972, Basteria, 36(2): 97-99.
Type Species. Xylopholas altenai Turner,
original designation.
i eneiied Characters. Valves and me-
soplax typical for the genus Xylophaga.
Animal long, not capable of retraction
within the FAlhios. 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-_
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. |
Wood-storing cecum large.
Remarks. This genus Gutters from Xylo-
phaga in having extended excurrent and
incurrent annals: contained in a common
sheath with siphonal plates at the posterior
extremity. These plates probably arose in- |
dependently but may be homologous with -
the siphonoplax of other pholads G. e., Pho- |
ladidea), which was carried posteriorly as
the animal elongated. However, the si- |
phonal retractor muscles of these species
insert on the valves rather than on the si-
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. Embryological studies are needed
to prove the affinities of the plates.
From Xyloredo, Xylopholas 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 decntOns. )
Xylopholas altenai Turner 1972
Plates 33, 34
Xylopholas altenai 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. Animal 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 Xylophaga. 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-
XYLOPHAGAINAE ° Turner 963
tinct except near the ventral margin,
slightly segmented, and with a small ven-
tral Ronee. Chondrophore and internal
ligament well developed. Posterior adduc-
tor muscle scar lar ge, 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 ee 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 dor eal 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
TABLE 20. MEASUREMENTS OF
XYLOPHOLAS ALTENAI.
Length Height
(mm) (mm) Location
1.8 1.8 holotype
1.0 1.0 Atlantique Sud, station 147
1.5 At Galathea, station 52
1.2 1.1 Gerda, station 66
1.9 1.5 Gerda, station 266
2.0 Rel Gerda, station 266
2.0 D5 Gerda, station 266
Obs O55 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 very acid.
However, the characteristic siphonal plates
readily identified the species. The two
specimens taken from a coconut husk
dredged off Sao Tomé 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,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
79°59'W) in 366 m; 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 Tomé 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 ane 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 pannel 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 differs
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 |
Xylophaga make a burrow more than five
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 caleareous tubes of Xyloredo but rath-_
er with the chimney of rock-boring pho-
lads, as in Parapholas Conrad Carmen
1955: 123).
The discovery of this teredolike 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 Hibee 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 nooi Turner
Plate 35
Xyloredo nooi Turner, 1972, Breviora, 397: 5-7, pls.
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-
XYLOPHAGAINAE ° Turner 265
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-
tracum. 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. Umbonal-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 muscles 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
TABLE 21. MEASUREMENTS OF XYLOREDO NOOI.
Length Height
(mm) (mm) Location
5.0 Sy, IL 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 very
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 calearious 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 larvae are
similar to those of Xylophaga. 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
|
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 mm.
from the anterior end, thus leaving room —
for boring activities and changing the di- |
rection of the burrow.
Range. Known 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, |
pls. 3-5 (from wood dredged by the Ingolf Expe-
dition at station 67, south of Eyrabakki, 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; Tumer, 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
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-
TABLE 22. MEASUREMENTS OF
XYLOREDO INGOLFIA.
Length Height
(mm) (mm) Location
PAS) 2.3 holotype
2.5 2.0 paratype
all 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. Autener 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 very 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 fives fourths length
of burrow. Tube relatively heavy, fatinked
with uniform, 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
thin, irregularly 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, aml in having the
valves longer than high. It differs from X.
nooi in having valves longer than high, in
having a isu rounded posterior slope, in
lacking the distinct groove on the inner sur-
face separating the valine from the posterior
slope, and in having the plates of the me-
soplax subrectangular and well calcified.
XYLOPHAGAINAE ° Turner 267
Range. Known only from the type local-
ity.
Specimens Examined. ICELAND: Ingolf Expedi-
tion, station 67, S of Eyrabakki (61°30'N, 22°30'W)
in 1,783 m.
Xyloredo naceli Turner
Plate 38
Xyloredo naceli Turner, 1972, Breviora, 397: 9-11, pl.
6, figs. 1-5 (USNCEL STU I-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 odes to July 1965). Holotype, MCZ 279638;
paratype, 279639.
Distinctive Characters. Burrow teredi-
nidlike, lined with a thin calcereous tube
marked with growth rings and covered
with periostracum. Shell Sailer to Xylo-
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
periostracum. Periostracal sheath covering
the animal between the valves and the
tube papillose.
Description. Shell globose, reaching 1.5
mm in length, thin, fragile, white, aah al
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. Umbonal—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
TABLE 23. MEASUREMENTS OF XYLOREDO NACELI.
Length Height
(mm) (mm) Location
1.1 2 holotype
1.0 1.0 paratype
1.2 1.3 paratype
1.5 1.5 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-
Ccareous, even in very young 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
ml/L, the pH was 7.84, and the hydrostatic
pressure 3000 psi (Muraoka, 1966b).
The embryonic valves still visible on
some of the specimens suggest that the
mature larvae are similar to those in Xy-
lophaga. Xylophaga muraokai 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 I-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 NO0014-91-J-1402,
Biological Studies on Marine Boring and
Fouling Mollusks.
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XYLOPHAGAINAE ° Turner 269
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Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
NS)
aI
i=
Plate 1. Xylophaga concava Knudsen from Pillsbury, station 526.
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 Daal
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.
i, Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
0.5mm
Plate 3. Xylophaga gerda Turner n. sp. from Pillsbury, station 944.
Figure 2. Inner view of left valve, showing posterior adductor muscle scar and low umbonal-— !
Figure 1. Outer view of right valve.
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. i
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. Dorsalview. 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 I/, station 124, showing the muscle scar and large number of ridges on the anterior slope of a specimen —
from hard wood.
XYLOPHAGAINAE ° Turner 275
0.5mm
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 /Ingo/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. |
~l
~]
XYLOPHAGAINAE ° Turner 2,
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 curvature 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 posterior dorsal margin, the umbonal reflection, and the inner surface of the mesoplax.
280 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
5mm
Plate 11. Xylophaga whoi 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 I8l1
Plate 12. Xylophaga profunda Turner n. sp. from U.S. Naval Oceanographic Office test site, Tongue of the Ocean,
Andros Island, Bahama Islands.
Figures 1-3. Holotype. Figure 1. Dorsal view showing round ridge posterior to the umbonal—ventral sulcus, the mesoplax, and
attached 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 posterior slope, as well as arrangement
and number of young. Figure 6. Enlarged young.
282 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
2
5mm
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
ales
1mm
Plate 15. Xylophaga abyssorum Dall.
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 Dall.
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, showing the mesoplax in place, standing off from the surface of the valves. Figure
2. Inner view of left valve of a paratype showing the 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 muraokai Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site |.
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 Museum of Comparative Zoology, Vol. 157, No. 4
1mm
Plate 19. Xylophaga muraokai Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site I.
Figure 1. Anterior view of hinge 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
3
1mm
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
Plate 21. Xylophaga washingtona Bartsch. |
Figures 1-3. Specimen from U.S. Naval Civil Engineering Laboratory Test Site |. 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
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.
bo
bo
9 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
5mm 5mm
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 showing siphons with lateral periostracal 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 -
f
of chondrophore.
XYLOPHAGAINAE ° Turner 293
15 hear: 16 6 17
Plate 24. Xylophaga depalmai 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
Plate 25. Xylophaga guineensis Knudsen.
Figures 1-3. Specimen from Pillsbury, station 260. Figure 1. Lateral view of entire animal showing 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
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, 2. Outer and inner views of left valve
Figure 3. Lateral view of entire specimen showing inflated mesoplax and siphons. Figure
Figures 5, 6. Outer and inner view of right valve of
Figures 1-4. Specimens from “replication reef,” Santa Monica Bay, California.
of specimen close to the holotype.
4. Dorsal view of entire specimen showing mesoplax and expanded foot.
the holotype (from Turner, 1955, pl. 90).
w
Ke)
~l
XYLOPHAGAINAE ° Turner
INNS
=
SS Sreoes
——_
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.
98 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
EAN
mameRe
ere
ee
: % ia
EEN
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 fringed 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 right 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
Gi i
ry igcs;) VAN WR eed <
\ my \\ uy ( \\\
i j (
Wa
( ) A ae
Ow
Plate 30. Xylophaga bayeri Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida.
Figure 1. Lateral view of holotype showing siphons and mesoplax in place. Figure 2. Dorsal view of holotype, showing 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
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.
XYLOPHAGAINAE ¢ Turner 301
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 11. Inner lateral view of mesoplax of a small specimen.
302 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
SSE Yo
Ki
N
LE
pape ys
\
Gs jist
Plate 33. Xylopholas altenai Turner from Gerda, station 66.
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 anteriorly so that the plates of the mesoplax are folded upward. Figure 7. Outer view of siphonal plate. Figure 8.
Inner view of siphonal plate.
Figures 1-3. Holotype.
XYLOPHAGAINAE ¢ Turner 303
1mm .
Plate 34. Xylopholas altenai Turner from Gerda, station 66.
Figure 1. Outer view of right valve showing 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
3
|
|
|
Plate 35. Xyloredo nooi 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
1mm
Plate 36. Xyloredo ingolfia Turner from Ingolf 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
Plate 37. Xyloredo ingolfia Turner from Ingolf 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 naceli Turner from U.S. Naval Civil Engineering Laboratory Test Site | (submersible test unit I-4).
Figure 1. Lateral view of holotype. Figure 2. Lateral view of specimen partially dissected from the wood showing the calcereous
tube with the anterior periostracal margin and the papillose periostracal sheath covering the animal between the tube and the
Figure 3. Anterior-lateral view of entire specimen showing foot and mesoplax. Figure 4. Inner view of left valve showing
valves.
Figure 5. Outer view of left valve.
muscle scars and chondrophore.
308
INDEX
Taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer t
species descriptions; t indicates table.
abyssorum, Xylophaga, 4, 12, 22, 23-24, 61-63
africana, Xylophaga, 4, 17-18
altenai, Xylopholas, 41-42, 80-81
atlantica, Xylophaga, 4, 27-28, 67
aurita, Xylophaga, 5, 31
bayeri n. sp., Xylophaga, 5, 12, 24, 38, 38-39, 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, Xylophaga, 4, 16-17, 51
growth series, 12
guineensis, Xylophaga, 5, 32, 34-35, 40, 72-73
hadalis, Xylophaga, 4
indica, Xylophaga, 5, 40
ingolfia, Xyloredo, 44, 44-45, 84
japonica, Xylophaga, 5, 24, 37, 39-40, 79
Lignopholas, 3
lobata, Xylophaga, 4, 22, 24
Martesia, 3
mesoplax
as character used in grouping, 3—5
variation in, 11-12
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
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, 45-46, 85 |
nooi, Xyloredo, 13, 43-44, 46, 82 |
obtusata, Xylophaga, 4, 20, 21
panamensis, Xylophaga, 4,18
Pholadidea, 40 |
pholadids, 3, 41 |
praestans, Xylophaga, By, Ie, Bul
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
tomlini, Xylophaga, 5
tubulata, Xylophaga, 4, 20, 21
turnerae, 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, Xylophaga, 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
=
aa
=
. =
. y
}
R ull etin oF ae
THE BOLAS SPIDERS OF
THE GENUS MASTOPHORA
ARANEAE: ARANEIDAE)
HERBERT W. LEVI
HERBERT W. LEVI
HARVARD UNIVERSITY VOLUME 157, NUMBER 5
CAMBRIDGE, MASSACHUSETTS, U.S.A. 5 February 2003
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
OR DISTRIBUTED BY THE
MUSEUM OF COMPARATIVE ZOOLOGY
HARVARD UNIVERSITY
Breviora 1952—
BULLETIN 1863—
Memotrrs 1865-1938
Jounsonia, Department of Mollusks, 1941-1974
OcCASIONAL PAPERS ON Mo.Luusks, 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 has been printed on acid-free permanent paper stock.
© The President and Fellows of Harvard College 2003.
Plate 1. Upper left, Mastophora diablo new species (Argentina, photo J. Abalos). Upper right, M. stowei new species (Kentucky,
photo K. F. Haynes and K. V. Yeargan). Middle left, MM. yeargani new species (Kentucky, photo K. F. Haynes and K. V. Yeargan).
Lower left, M. alvareztoroi |barra and Jiménez new species (Texas, photo M. Stowe). Lower right, M. fasciata Reimoser (Costa
Rica, photo W. Eberhard).
THE BOLAS SPIDERS OF THE GENUS MASTOPHORA
(ARANEAE: ARANEIDAE)
HERBERT W. LEVI"
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 known
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.
Agatostichus is synonymized with Mastophora. With-
in this paper, Ibarra and Jiménez describe a new spe-
‘cies from Chiapas and Texas. Evidence from palpal
morphology indicates a relationship of Kaira with
Taczanowskia 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 Euglyptila is synonymized with
Ordgarius.
INTRODUCTION
Female bolas spiders spend the 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 moths 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 pheromone
was found by Eberhard (1977), and chem-
ical analyses were conducted by Stowe et
al. (1987) and Gemeno et al. (2000). An
approaching male moth 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.
Bull. Mus. Comp.
las, hurled at the moth. Moths stick to the
bolas, whereas detachable wing scales per-
mit moths to escape from most orb webs.
The unusual behavior of Mastophora, first
observed 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 made. 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 from 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-
Zool., 157(5): 309-382, February, 2003 309
310
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 Cladomelea 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. phrynosoma 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
meriateccrn flies, primarily Psychodidae
(moth midges), oven 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 (Bréthes, 1909)
and M. dizzydeani produced 826 eggs
(Eberhard, 1981). Oviposition of M.
hutchinsoni (a North American species)
takes place in fall and spiderlings emerge
in May (Yeargan, 1988). This differs for M.
cornigera. Glitch sizes for M. hutchinsoni
ranged from 178 to 275. The sex ratio ap-
proached one to one (an unusual excep-
tion was observed in one egg case of M.
phrynosoma, 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 and
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 cryptic
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 (Ebem
haved 1981). A summary of research was_
reported by Yeargan (1994).
METHODS AND ACKNOWLEDGMENTS
The collections of the following institu-
tions and individuals were used.
American Museum of Natural]
History, New York (N. Platnick, |
L. Sorkin)
AMNH
BMNH
ECOTAR
FCMU
FMLT
FSCA
BSP
INHS
IRSNP
MCN
Natural History 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,
le v5 JE Jee rexchti))
Florida State Collection of Ar-
thropods, Gainesville, Florida
(G. B. Edwards)
Instituto Butantan, Sao Paulo,
Brazil (A. Brescovit)
Illinois Natural History Survey,
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 Argentino de Ciencias
Naturales, Buenos Aires, Ar-
gentina (M. E. Galiano, C. L.
Scioscia)
Museu de Ciéncias Naturais,
MCP
MCZ
MKS
ML]
MLP
MNHN
MNRJ
MUSM
MZAQ
MZSP
NHMW
NMB
NMP
OSU
QMB
TAMU
MASTOPHORA ® Levi 311
Fundagao Zoobotanica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil (E.
H. Buckup, M. A. L. Marques)
Museu de Ciéncias, Pontificia
Universidade Catélica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil (A.
A. Lise)
Museum of Comparative Zo-
ology, Cambridge, Massachu-
setts
M. K. Stowe, Gainesville, Flor-
ida
M. L. Jiménez, La Paz, Mexico
Museo de Universidad Nacion-
al, La Plata, Argentina (C.
Ituarte, L. A. Pereira)
Muséum National d’ Histoire
Naturelle, Paris, France (C.
Rollard)
Museu Nacional, Rio de Janei-
ro, Brazil (A. B. Kury)
Museo de Historia Natural,
Universidad Nacional Mayor
de San Marcos, Lima, Peru
(Diana Silva D.)
Museu, Departamento de
Zoologia da Escola Superior de
Agricultura “Luis de Queiroz,”
Piraciba, SA0 Paulo State, Bra-
zil (G. J. de Moraes)
Museu de Zoologia, Universi-
dade de Sao Paulo, Sao Paulo,
Sao Paulo, Brazil (E. M. Can-
cello, R. Pinto da Rocha)
Naturhistorisches Museum, Vi-
enna, Austria (J. Gruber)
Naturhistorisches Musuem,
Basel, Switzerland (A. Hinggi)
Natal Museum, Pietermaritz-
burg, South Africa (D. A. Bar-
raclough, C. Conway)
Ohio State University, Marion,
Ohio (R. A. Bradley)
Queensland Museum, Bris-
bane, Queensland, Australia
(R. J. Raven)
Texas A&M University, College
312
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)
USNM
ZMUC
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. Kury. 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 laboratory. 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. Thue N. and M. L. Jiménez
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 Wetmor ev 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
small and difficult genitalia of 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-
ers 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 Hoyer’s 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 “ilies ated 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 warp sclerites. —
Gertsch (1955) in his revision did not’
interpret the two original descriptions by.
Banks (1898) of Ordgarius obesus and O.
corpulenta. 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 Marx died. Banks (1898) pointeds
out Le the time:
> Hoyer’s 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 bottle.
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 Hibercles 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 humps
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 leastn two instars
to mature. The males of only a few species
are known, because only ‘W. Eberhard,
Mark Kk. Stowe, and Ken V. Yeargan have
raised individuals. The males of most
southern South American species remain
MASTOPHORA ° Levi 3138
unknown. The palpi of different species
are surprisingly similar and the determi-
nation of males presents a challenge.
Immatures coming out of the egg sac
have visible median’ Raiborcles 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
detenumed 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.
Gonsch (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
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-
melea species (Fig. 443). The different
species of Ordgarius and Cladomelea show
greater morphological differences from
each other than do those of Mastophora.
Different American 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 ( Sedna 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-
tanéo-hémolytique gangrénieux” 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. es-
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
(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 Cladomelea
Simon, 1895 (Figs. 434-444). Both Ord-
garius and Cladomelea species handle the
Boles with the second leg and, unlike Mas-
tophora, which swings the bole in a pen-
dulumlike motion, they whirl the bolas,
Ordgarius when a moth approaches, ane
Gindomeion 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 Euglyptila
Simon, 1908, from northern Vietnam
(Tonkin). Euel yptila is synonymized below
with O 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, Cladomelea
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 Cladomelea 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 |
Yeargan (1994) studied the relationships
with other genera. Scharff and Coddington
° Roewer (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 Stand (Neave, 1939a: 9: Bon- |
net, 1955: 124).
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, Cyrtarachne,
Poecilopachys, and Pasilobus; Table 1).
Robinson and Robinson (1975) first sug-
gested that the web of Pasilobus 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-
apomorphies, 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 syn-
apomorphy of Kaira, Metepeira, and Tac-
zanowskia. One of the synapomorphies 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-
paida, Eriophora, Ocrepeira, Acacesia,
Cyrtophora, and many others). The shape
of the median apophysis of the palpus of
Pasilobus 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
MASTOPHORA ® Levi 315
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.
cornigerum Hentz designated by Simon. Neave,
1939b: 484. The gender of the name is neuter.
Bonnet, 1957: 1995. First synonymized by Bréthes,
1909.
Agatostichus Simon, 1895: 885. Type species A. leu-
cacantha Simon by original designation and mon-
otypy. Neave, 1939a: 86. Gertsch, 1955: 250. NEW
SYNONYMY.
Agathostichus:—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. leucabulba
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. alvarextoroi, and also not for the type
species. Tubercles also are found between
eyes in M. corpulenta (Figs. 336, 337).
A revision of all species of the genus was
made by Mello-Leitaéo (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-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
316
‘UOHRAIOSGO OU “YURI ‘JUESqR IayRIRYO *— squosoid ToyORIeypo “4 %
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317
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psourdsiq
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snowuofng
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318
lication of the two 1931 articles, but Mel-
lo-Leitao’s must have been published 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
Cladomelea and Ordgarius, which may
have horns (Fig. 423), by using the first
pair of legs to Thence the bole: Clado-
melea and Ordgarius use the second pairs.
Description. Female. Carapace color
variable, often uniform orange to strongly
marked. Legs usually not Ramded! Apdo
men often donee anteriorly, lighter pos-
teriorly, venter often with a edian white
square or rectangle (black in M. hutchin-
soni and some M. phrynosoma). Carapace
about as wide as long. Eyes small, sub-
equal. Anterior eta eyes slightly larger,
lateral eyes smallest. Median ocular tr ap-
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 mcd eye Giammcters: Carapace
very 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. bisaccata 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
or setose. Some species (M. alvareztoroi,
M. felis, and M. haywardi) 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 Ge about 1.1 fines 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, escomeli, obtusa, felis,
holmbergi, reimoseri, satan, and diablo).
Their carapace is high with vertical sides
(Fig. 408); the sides OE the thorax have tu-
bercles (Fig. 409), with short, white setae
between; ane the abdomen may have high,
tube-shaped horns (Fig. 413). The Albee
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 (arafia cabeza de gato)
because of their resemblance to the head
of a cat, as can be seen in Figure 421.
‘S4 VA Roe ONO
UE MEXICO A
NX
“—} BELIZE
Uw
HON.
GUATEMALA-S 4 = / 5
SICA
a
COSTA RIGA
- Wz
PANAMA +3 ap
we
JAMAICA
MASTOPHORA ® Levi 319
rare TRINIDAD
f n= ~
VENEZUELA SOn
aN’ A
SURINAM
oy €oLomeia
= / NS
fe
>- ECUADOR
GALAPAGOS
Map 1.
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. bisaccata 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.
Sk :
NS 2227 YRUGUAY
Approximate number of species in various American regions.
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
IL Abdomen without humps (Figs. 10, 24,
88, 130); swellings may be visible in
pronler (igs ys) eee ee eee 2
= Abdomen with distinct humps or tuber-
cles (Figs. 137, 145, 298, 331, 401) _. 15
(1). INoxrthyAmericas(Mapi2) e= serene eee 3
= South America (Mapsi3 44) oa 11
3(2) Abdomen subtriangular, with anterior
320
(2):
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
SH] B) )) eae ee elt fee sui isa) SUE Ny phrynosoma
Abdomen without anterior lateral swell-
ings (Figs. 10, 17, 24); epigynum with-
out posterior lips (except apalachicola)
BE aoe te ete ne eer Pe Ne Ne ay Se ee 4
Posterior of epigynum with slits framed
by lateral lips (Fig. 20); southeastern
United States (Map 2C) ___ apalachicola
Slits without lips (Figs. 13, 27) 5
Slits short, their length distant from ven-
tral margin (Fig. 80); eastern United
States (Map 2A) yeargani
Slits approaching close to ventral margin
Geass 7), Ge 6
Dorsum of abdomen with bisaccata pat-
tern, anterior orange to gray with
spaces and lines (Figs. 42, 53, 65)
Abdomen marked otherwise (Figs. 10,
24)
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
(Eig SrA ai S)) ee a eer 8
Slits 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
2G) stowei
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,
Oye lNeratoey Se ee ee 10
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)
ey nicenee ie, tek Na SE wee eee felda
Abdomen with only anterior median
area black (Fig. 10); atria at level with
large seminal receptacles, seminal re-
ceptacles less than their diameter dis-
tant from ventral border (Fig. 14);
lmlloyato eV (IM eho) XC) satsuma
Abdomen hirsute (Figs. 109, 110), slits
and atria separating ventrally (Figs.
112, 113); Tucuman, Argentina (Map
3C) haywardi
Abdomen with few setae (Figs. 102,
116): atria otherwise (105, 119)
12(11). Abdomen with many black spots and
dark lines (Figs. 130, 131): southern
Brazil (Map 3B) carpogastra
16(15).
17(16).
18(17).
Abdomen marked otherwise (Figs. 102,
116, 123)
. Abdomen with light longitudinal lines
(Figs. 102, 103); atria approaching
each other ventrally (Fig. 105); Santa
Catarina, southern Brazil (Map 3C) —
a catarina
Abdomen marked otherwise (Figs. 116,
]ED 23) ea A ee NU Leelee ccaeelt ae ae eee es 14
. Abdomen marked with pair of black
rings, open anteriorly (Fig. 116); Sao
Paulo, Brazil (Map 3C) _.._ corumbatai
Abdomen with two black discs (Fig.
123); Lara, Venezuela (Map 3A) __.... lara
Abdomen with many dorsal or lateral
humps (Figs. 267, 298, 300, 310, 331,
RSIS) WR Mer ete Tania tee Tae ee ee URE Sa 16
Abdomen with one pair of humps (Figs.
155, 171), rarely median area of ab-
domen swollen (Fig. 325) 19
Humps anterior on each side of abdo-
men (Fig. 267); Santiago del Estero,
Acgentunas (Mayor Si) ee eae abalosi
Humps or tubercles dorsally (Figs. 298,
MOL SSM Brae Nike cle aes aaa 17
Carapace with short tubercles (Figs. 329,
330); abdomen with numerous small
dorsal tubercles (Figs. 331, 332):
northern Argentina to Buenos Aires
Province (Mapi3G@)ss sans conifera
Carapace with large tubercles (Figs. 297,
3()9))gesie 2 ane Can eT UBS aera pseu eaeee 18
Carapace with median tubercles spine-
shaped (Fig. 309); Panama (Map 2F)
LS Sn SEED eh in TS soberiana
Carapace with median tubercles cone-
shaped (Fig. 297); Texas to Chiapas
(Map 2F) alvareztoroi
; Carapace median tubercles same size or
longer than horns (Figs. 288, 316, 323)
AS Sata et lichen AA te et hed i la ln 5 EN 20
Carapace with median tubercles smaller
thanshoms)(Higs: 1365.38)l)) )iemseene 22,
. Median tubercles spine-shaped, very
long (Fig. 316); Bahia to Rio de Ja-
neiro, Brazil (Map 3G) _....... leucacantha
Median tubercles otherwise (Figs. 288,
SOS) es OE NR EE AE Sea ee 21
. Median tubercles cone-shaped (Fig.
323); Rio Grande do Sul, Brazil (Map
3G) brescoviti
Carapace tubercles large, rounded on
dark carapace (Fig. 288); Texas, Mex-
Itcfo) (UN eto) PAN) Se leucabulba
. Carapace with sides vertical (Figs. 366,
380), sides with tubercles (Figs. 367,
381); Mexico to South America (Map
721) nie ee eel ae TEs me ETE Sune ta 39
Carapace with sides slanting (Figs. 135,
169): sides with few or no tubercles
(HigstllS65ell7(0;"Miapse2 3) ea=eeaenes 23
MASTOPHORA ° Levi
apalachicolag@
satsuma YW
stowei @
A felda
g@ timuqua
@ yeargani
archeri o
hutchinsoni @
alachuaa YW
cornigera @
seminole A
vaquera
SP.
Ss
Qq
Se ;
—>
@ alvareztoroi
wv corpulenta ;
A fasciata au
@ Jeucabulba ° §) /\
@ soberiana ‘y iG
Map 2. Distribution of Mastophora species of North and Central America.
321
322) Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
catarina @
corumbatai V
hayward; 4&
@ dizzydeani
@ fasciata
BRAZIL - ~~~ - A BRAZIL~ { BRAZIL_ |
LZ - Tbe =e)
NN MS «\ Woe cw
KN Q
abalosi Vv
comma Vv
melloleitaoi @
yacare A
ypiranga
cranion A
longiceps w }
pesqueiro @
pickeli A.
piras
extraordinaria
F
brescoviti V
conifera @
leucacantha @
Map 3. Distribution of Mastophora species of South America.
34(33).
. North America and Cuba (Map 2) _....... 94
Central and South America (Map 3)
. Cuba (Map 2G); abdomen with anterior,
lateral swellings (Figs. 145, 146)
ee ee eee ee vaquera
Continental North America; abdomen
without lateral swellings (Figs. 137,
Ay Sy) Re nem Aik ey otic se Ree eee 25
. Posterior of epigynum with slits and atria
approaching each other ventrally
(Figs. 159, 160); northeastern United
Steltes (Mizyo 2B) ee hutchinsoni
Slits and atria otherwise (Figs. 140, 174,
ARS SS) prgetrrere mee eccrine a Maas oases EA 26
. Posterior of epigynum with slits framed
by a lip on each side (Fig. 188); south-
eastern United States (Map 2E) — archeri
Slits without lips (Figs. 140, 174) if
. Slits parallel and approach ventral bor-
der (Fig. 174); eastern United States
to California and Honduras (Map 2G)
po ie Eee SESE Ebon wed Tecate ok Aaah ae cornigera
Slits short, their length apart from ven-
tral border (Fig. 140); southern Flori-
(6 (Fie TT cy cee ARE Se aR ae seminole
. Central America and northern South
Niraveraver) (we) leteinby =e 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) —
oe a ee fasciata
Humps of abdomen narrow (Fig. 207);
slits separating ventrally (Fig. 210);
Colombia to Peru (Map 3A) — dizzydeani
. Humps of abdomen extended to a point
(Figs. 271, 272); Santiago del Estero,
Argentina (Map 3E) comma
BOERS Sa pe Vary a aL oe pg ee ee Hee Ve et 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
(Nia c3ID)) i lee eee pesqueiro
Abdomen without such marks _ 32
. Posterior of epigynum with a lip on each
side between slits and ventral margin
(Fig. 263); Uruguay (Map 3E) _.. yacare
Posterior of epigynum without such lips
(ale OG) ae ee 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(34).
36(34).
45(44).
MASTOPHORA ® Levi 323
Epigynal slits and atria approaching each
other ventrally (Figs. 256, 286)
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
GAY C0) S6)) D)) te eee lee eee Zales piras
Anterior of abdomen light, light area
bordered posteriorly by dark (Fig.
283); epigynum as in Fig. 286; south-
erm Brazil, northern Argentina (Map
3 Fg ete LA trac de el extraordinaria
Anterior of abdomen black (Figs. 218,
253, 276)
. 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
. Atria with median knobs facing each oth-
er (Fig. 280); Parana to Buenos Aires,
Argentina (Map 3E) melloleitaoi
Atria without knobs (Fig. 222); Pernam-
buco; Brazil (Map 3D) = pickeli
. Central America (Map 4A); posterior of
epigynum with dark patch dorsally in
each of pair of depressions (Fig. 341)
corpulenta
OUnNerraelo ns 40
. Galapagos (Map 4A); abdomen with only
tiny humps (Fig. 346); epigynum slits
each with a loop ventrally (Figs. 348,
349) rabida
Other regions; abdomen with larger
humps
. Peru (Map 4A); posterior of epigynum,
between slits, with median area having
a bulge on each side (Fig. 355) escomeli
Otherre gions yeast arsme een wen Lt 42
. Chile (Map 4B); posterior of epigynum
with a dark spot dorsally in each ad-
jacent depression (Fig. 415)
Eh a ee Na gasteracanthoides
Brazil to Argentina
2). Abdomen with humps placed on swollen
area (Figs. 364, 365); Pernambuco
State, Brazil (Map 4C) obtusa
Humps not placed on swollen area (Figs.
368, 369)
. Dark patch (atria) placed dorsal or mid-
dle of seminal receptacles (Figs. 392,
404)
Atria absent or placed ventrally of sem-
inal receptacles (Figs. 372, 379, 386)
tee fy dhe Dalal Uae ead dicen seat k eS 46
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
pura?”
: CB) canaGua
B corpulenta
@ escomeli
w ‘abida
wv felis
& holmbergi
QO obtusa
@ ‘eimoseri
diablo
D
Map 4. Distribution of Mastophora species of the M. gasteracanthoides group.
= Atria lateral, placed outside of depres- = Slits without such bend (Figs. 371, 378)
sionu(Higs 404) mortherneicentralAve? Tay) | eee to ae ee ee eee AT
gentinan(Mapw4 1D) ese) ser diablo 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 AG) 18s e ete eer oe ee reimoseri sievalll (UMleie) 4G) ee felis
Slits parallel (Fig. 378); Paraguay, San-
tiago del Estero (Map 4C) __.
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)
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)
Terminal apophysis shorter than embo-
lus (Fig. 48); median apophysis of pal-
pus very short (Figs. 48-50); Florida
(Mian? CG) aes eee ee
Terminal apophysis as long or longer
than embolus (Figs. 7, 60)
Base of embolus large (Figs. 60, 61);
eastern United States (Map 2B) ———
holmbergi
alachua
ae ee Se ea SI a bisaccata
Base of embolus small (Figs. 72, 73)
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) _....
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
Wnited States! (Map. 2©) = es
Terminal apophysis almost as long as
embolus (Figs. 94, 177, 180)
Terminal apophysis about half length or
less of embolus (Figs. 83, 191) ...........
Base of median apophysis longer than
wide (Figs. 177, 180); southern United
States to California and Central Amer-
jean (Mawr 2 Chon. sn: SA eee
Base of median apophysis short (Figs.
94, 98); eastern United States (Map
2D)
Base of median apophysis large, touch-
ing embolus (Figs. 83, 84); eastern
United States (Map 2A)
Base of median apophysis small (Figs.
37, 191)
Length of narrow part of median apoph-
ysis as wide as base in ectal view (Fig.
193); Gulf Coast, Kansas (Map 2E) —
4
hutchinsoni
stowei
cornigera
phrynosoma
yeargant
Ba nee ae wR AEN ce pe Ae RP archeri
Length of narrow part of median apoph-
ysis longer than width of base (Fig.
39); Virginia to North Carolina (Map
CIN) eviee Sree e ine Mier ikea Ae!
timuqua
MASTOPHORA ° Levi 325
Kaira altiventer (O. P.-Cambridge), new
combination
Epeiroides fasciolata O. P.-Cambridge, 1889: 15, pl.
8, fig. 5, 3d. Male from Bugaba, Panama, in
BMNH, examined; now lost. Keyserling, 1893: 309,
pl. 16, fig. 228, d. Male from Guatemala.
Kaira altiventer O. P.-Cambridge, 1889: 56, pl. 3, fig.
13, 2. Female from Veragua |Veraguas Prov.], Pan-
ama, in BMNH, examined. Levi, 1993b, 213, figs.
3-22, 2, 6. NEW SYNONYM Y.
Aranea fasciolata: —F. P.-Cambridge, 1904: 519, pl.
51, fig. 5, d. 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. P.-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. Abdomen
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
Carapace teat mm long, 4.0 wide in tho-
racic region, 2.4 wide a 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 ccminal 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), ariel Alee. by the larger
depression in ras 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 ae 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 very 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
wide at lateral eyes. First femur 3.3 mm,
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 Boreal 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-
em Florida (Map 2C).
Paratypes. 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,
12 (M. K. Stowe 2116, FSCA). Levy Co.: Manatee
Springs State Park, 10 Nov. 1992, 12 (M. K. Stowe
2114, MCZ).
Mastophora felda new species
Figures 22—28; Map 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 Stemi and legs. Dis-
tal leg spetales dark orange- esate Abdo-
men anterior, sides, and enter gray (Fig.
24), center and posterior whitish; venter
MASTOPHORA ® Levi 327
seminal receptacle
now atrium satsuma
terminal apophysis
radix embolus
tegulum conductor area
median apophysis
Figures 1-7. Mastophora. 1-4, female. 1, M. bisaccata, left tip of chelicera 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. MM. 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. WM. 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
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.
Be
Distribution.
(Map 2A).
South-central Florida
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, digs leg articles with dark
brown rings. omens 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
without humps. Total length 8.5 mm. Car-
apace 3.5 mm long, 3.4 wide in thoracic
region, 1.8 wide at lates al eyes. First femur
2.9 mm, patella and tibia 4.1, metatarsus
2.7, tarsus 0.8. Second patella and tibia 3.3
mim, 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
ies 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. cornigera by the prox-
imal position of the radix (Fig. Sas
The egg sac is shown in Figure 445.
Distribution. North C@xolina to north-
ern Florida (Map 2A).
Paratypes. NORTH CAROLINA Moore Co.:
reared from egg sac in spring 1941, 3d (J. Perry, M.
K. Stowe 2113, “MCZ). FLORIDA Alachua Co.: Deve
ils Millhopper State Park, reared spring 1992, ¢ al-
lotype, 11 paratypes (M. K. Stowe 2101, MCZ,
AMNH); spring 1992, raised 3d (M. K. Stowe 2106,
FSCA). Levy Co.: Manatee Springs State Park, 5 Nov.
1987, 12 (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
timuqua
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, ectal.
Figures 40-50. WM. 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
Mastophora alachua new species
Figures 40-50; Map 2G
Holotype. Female holotype from Devil’s Millhopper
State Park, Gainesville, Alachua Co., Florida, 19
Nov. 1983 (Mark K. Stowe 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, Sv ide
in thoracic region, 2.2 wide at ieteral 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 oh
pair of adjacent humps. Total length 1.6
mm. Carapace 0.78 mm long, 0.67 wide in
thoracic region, 0.52 wide Ae lateral eyes.
First fom 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.
Diagnosis. Mastophora alachua is distin-
guished by the abdomen, slightly triangu-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
lar in shape and having markings like those
of M. bisaccata (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.
AE)
Paratypes. FLORIDA Alachua Co.: Devil's Mill-
hopper State Park, 21 Nov. 1983, 12 (M. K. Stowe
106, MCZ); no date, prob. 1984, allotype d (M. K.
Su 2102, MCZ); Gainesville, 1 Nov. 1990, 12 (M
. Stowe 2115, FSCA).
Northern Florida (Map
Mastophora bisaccata (Emerton)
Figures 51-62, 446; Map 2B
Cyrtarachne bisaccata Emerton, 1884: 325, pl. 34,
fig. 11, °, pl. 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, pl. 2, fig.
35, 2. McCook, 1894: 198, pl. 12) figs. 2: 3, oe
Glyptocranium bisaccatum:—Bonnet, 1957: 1996.
Ordgarius obesus Banks, 1898: 250, pl. 15, fig. 9, °.
Two female syntypes from La Chuparosa [Chupar-
rosa, San Luis Potosi], Mexico, in CAS, destroyed.
NEW SYNONYMY.
Mastophora_ bisaccata:—Mello-Leitaéo, 1931: 71.
Roewer, 1942: 900. Kaston, 1948: 232, figs. 737—
740. Gertsch, 1955: 242, pls. 3-5, pl. 6, figs. 1, 4;
text figs. 19-23, 35, 43, 44, 2, ¢d. Platnick, 1997:
513. Platnick, 2001.
Note. Atkinson did not tell how Cyrtar-
achne multilineata differs from bisaccata,
although he mentioned the latter species.
The large size, total length 11 and 13 mm,
abdomen 13 and 15 mm wide, suggest ne
had M. bisaccata. The name was first syn-
onymized by Banks (1910).
Ordgarius obesus differs from bisaccata,
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. bisaccata and the
carapace, unlike most Mastophora, is trun-
cate. There is no doubt that this was M.
MASTOPHORA ° Levi 331
Se
Wo stowei
bisaccata
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 chelicerae. 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
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 humps (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 with white line in middle, branch-
ing posteriorly into tubercles (Fig. 6). Ster-
num white. Lateral eyes smaller than me-
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 were made from
the female holotype and males from Ar-
kansas and Ohio. An egg sac collected with
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 cemetery 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, 12 (AMNH).
NEW YORK Nassau Co.: Long Island: Sea Cliff, 1d
(MCZ). NEW JERSEY Elizabeth Co.: Roselle Park,
25 Sep. 1910, 12 (AMNH). Middlesex Co.: New
Brunswick, July 1930, 12 (AMNH). PENNSYLVA-
NIA Westmoreland Co.: 4.8 km S Rector, 13 Sep.
1966, 12 (B. Vogel, DMNS). OHIO Jackson Co.:
Oak Hill, 12 (R. A. Reller, OSU). Logan Co.: Can-
twell Cliffs, 8 Sep. 1935, 12 (OSU). Butler Co.:
Bachelor Woods, Oxford, 11 July 1998, 1d (D. M.
Golden, OSU). DISTRICT OF COLUMBIA Wash-
ington, Sept., 1d (Fox, CUC, AMNH); summer
1935, 12 (H. E. Ewing, USNM). VIRGINIA Falls
Church, 12, 1¢ (MCZ). Powhatan Co.: Powhatan,
Sep. 1984, 12 (A. Moreton, MCZ); 1985, 12 (A.
Moreton, MKS). KENTUCKY Jessamine Co.: imm.,
73 raised spring 1995 (K. V. Yeargan, KVY). SOUTH
CAROLINA Oconee Co.: Clemson College, 1°
(MCZ). Lexington Co.: Batesburg, 12 (MCZ).
GEORGIA Fulton Co.: Atlanta, 2 Aug. 1937, 12 (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. MM. 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, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
334
W. Fattig, AMNH). FLORIDA Alachua Co.: Devil's
Millhopper State Park, July, Aug. 1978-1981, 52 (M
K. Stowe, MKS); June, July 1981, OMEN Ke Stowe,
MKS): 19 Nov. 1983, 12; Gainesville, 26 July 1980,
penult. (C. Hieber, MCZ); High Springs, Oct. 1935,
12 (H. H. Simpson, USNM). Indian River Co.: Se-
bastian, Apr. 1944, 12 (G. Nelson, MCZ). Lake Co.:
Umatilla, Sep. 1954, 12 (M. W. Tyler, AMNH). Sum-
ter Co.: Bushnell, 24 Oct. 1979, 12 (W. Edwards,
FSCA). ILLINOIS Franklin Co.: Zeigler, 10 May
1928, 12 (J. K. Carlovic, MCZ). Jackson Co.: Little
Grand Canyon, S. Murphysboro, 5 Sep. 1971, 1d (N.
Magnuson, JAB). Williamson Co.: Canterville, field of
cemetery, 14 Oct. 1978, 12 (R. Reith, JAB). MIS-
SOURI St. Louis Co.: 17 Feb. 1940, 1d paratypes of
M. archeri (W. M. Gordon, AMNH). ARKANSAS
Carroll Co.: Berryville, July 1942, 1d (C. Wilton).
MISSISSIPPI Harrison Co.: Gulfport, 12 (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°3 6.0'N, 82°22.0'W, 7 Dec. 1987 (M. Stowe
07001), in MCZ. The species is named after the
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-
most 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
eiael milo) S45 saonen, anixel BOs amon B2.
Length of first patella and tibia 1.3 times
width of carapace.
Male allotype. Carapace gray-orange
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 were 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. bisaccata by being small-
er in size and having the atria of the epi-
gynum ventrally departing from each other
Fi igs. 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.
Dicom Widespread in the eastern
United States (Map 2C).
Paratypes. CONNECTICUT Hartford Co.: Rain-
bow nr. Windsor, 9 Aug. 1939, 12 (A. de Caprio,
USNM). OHIO Logan Co.: Old Man’s Cave, 12 Sep.
1924, 15 (OSU). VIRGINIA Arlington Co.: Arling-
ton) 7 Septyl953) 121k Vv; Ker Omiein AMNH). IL-
LINOIS Pope Co.: Dixon Spring State Park, 7 Sept.
1974, 12 (J. A. Beatty, JAB). KENTUCKY Fayette
Co.: Lexington, Oct. 1997, 1°, egg sac (K. V. Yeargan,
KVY); 6 Aug. 1998, 14d (K. Yeargan, KVY):; Cold
Stream farm fence, 16 Oct. 1998, 32 (K. Yeargan,
KVY). Jessamine Co.: 3 Aug. 1995, 75 (K. Yeargan,
KVY). NORTH CAROLINA Haywood Co.: Canton,
12 (Holden, MCZ). GEORGIA 12 (MNHWN 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 chelicerae. 100, frontal. 101, lateral. 102,
103, carapace and abdomen. 102, dorsal. 103, lateral. 104-106, epigynum. 104, ventral. 105, posterior. 106, posterior, cleared.
Figures 107-113. MM. haywardi Birabén, female. 107, 108, carapace and chelicerae. 107, frontal. 108, lateral. 109, 110, carapace
and abdomen. 109, dorsal. 110, lateral. 111-113, epigynum. 111, ventral. 112, posterior. 113, posterior, cleared.
MASTOPHORA ° Levi 335
Figures 114-120. MM. 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
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,
ES X(Se1G-aWellsos ke):
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 white marks anteriorly (Fig. 77),
posterior white, 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,
ALS) ride 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. Sternum orange. Cox-
ae and distal leg articles orange. Abdomen
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
oo
sac of female M. yeargani (by K. Yeargan).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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. bisaccata and M. stow-
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. bisaccata (Fig.
62) by having a longer median apophysis
(Fig. 85) and om M. cornigera (Figs. 177
1 79) by having only a short terminal
apophysis in the palpus (Figs. 82, 83), and
by having the median apophysis Ol anait=
ferent shape, and the embolus wider (Figs.
Soma):
Distribution. From New York to Ken-
tucky (Fig. 448).
Paratypes. NEW YORK nr. New York City, on
Amalanchier sp., 12 (AMNH). KENTUCKY Mercer
Co.: Feb. 1995 egg sacs, 29 Sept. 1995, 7 imm., 4d
(K. V. Yeargan, KVY). Garrard Co.: Feb. 1995, egg
sacs, Sept. 1995, 6 imm., 16d (K. V. Yeargan, KVY).
Mastophora phrynosoma Gertsch
Figures 86-99, 449, 450; Map 2D
Mastophora phrynosoma Gertsch, 1955: 245; pl. 6,
fig. 5, text figs. 24-27, 31, 2. 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.
AGdewen anter iorly 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
7
nor 5
ee ~
SF
carpogastra
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
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 ies 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, aval 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 phrynosoma is
distinguished by the subtriangular shape of
the Aidomien. 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 wider
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 (Arundinaria gi-
gantea) at night at 0200 h. Males were col-
lected crawling on a table in the labora-
tory, 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,
observed 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.
(Map 2D).
Eastern United States
Specimens Examined. CONNECTICUT New Ha-
ven Co.: Mount Carmel, 4 Sept. 1946, 12 (K. M.
Somerman, INHS 4748). Meriden Co.: South Meri-
den, Oct. 1945, 12 (H. L. Johnson, USNM). NEW
YORK New York City, on Prunus, 12 (AMNH).
OHIO Cuyahoga Co.: Sagmore Picnic, Sagmore
Hills, Buckeye Trail area, 16 Sept. 1999, 12 (K. Brad-
ley, OSU). MARYLAND Anne Arundel Co.: Annap-
olis, 21 Sep. 1941, 12 (M. H. Muma, AMNH). How-
ard Co.; Colombia, Snowden River Parkway, 26 Sep.
1994, 12 (M. Harden, USNM). Montgomery Co.:
Cabin John, 10 Nov. 1943, egg sacs (I. N. Hoffman,
USNM). VIRGINIA Falls Church, 22 (N. Banks,
MCZ). KENTUCKY Fayette Co.: Lexington, Raven
Run, 1989, 19? (K. V. Yeargan, KVY); 17 May 1999,
egg sacs with only dd (K. V. Yeargan, KVY, MCZ).
GEORGIA Fulton Co.: Atlanta, 21 Aug. 1944, 12 (F.
W. Fattig, AMNH). FLORIDA Alachua Co.: Gaines-
ville, Apr. 1988, 1¢ (M. K. Stowe, MKS); Devil's
Millhopper State Park, 15 July 1980, 12 (M. K.
Stowe, MKS); spring 1992, 2d (M. K. Stowe, MKS):
spring 1994, 8d (M. K. Stowe, MKS). ALABAMA
Monroe Co.: Randon’s Creek, 19 Oct. 1941, 12 (A.
F. Archer, AMNH). INDIANA 12 (A. Petrunkevitch,
NHMW). ILLINOIS Jackson Co.: Carbondale, in
woods at night, 4 Aug. 1967, 12 (J. M. Nelson,
AMNH). MISSOURI Wayne Co.: Markham Spring,
Mark Twain National Forest, 14 Oct. 2000, 12 (E
L. Quinter, AMNH). St. Louis Co.: 17 July 1940, 13
paratype of M. archeri (W. M. Gordon, AMNH).
TEXAS Walker Co.: Huntsville State Park, on elm
bush, 27 Sep. 1987, 12 (W. R. Martin, TAMU).
Mastophora catarina new species
Figures 100-106; Map 3C
Holotype. Female from Pinhal, Est. Santa Catarina,
Brazil, Dec. 1948 to Jan. 1949 (A. Maller), 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 brown. 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
wie in thoracic region, 1.8 wide at tae al
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 Birabén
Figures 107-113; Map 3C
Mastophora haywardi Birabén, 1946: 327, figs. 1-3,
2. Female holotype, from Tucuman, Argentina, in
MASTOPHORA ® Levi 339
MLP. examined.
2001.
Brignoli, 1983: 274. Platnick,
Description. Female holotype in poor
condition. Carapace brown, with a lighter
square area in thoracic region (Fig. 109),
and with long white setae. Stemi lighter,
Legs frown Abdomen brown (the Thole.
type has an injury, a large transverse gash
anteriorly), posterior and: sides darker gray
than anterior (Fig. 109), with some long
white setae and indistinct humps ate
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 formate 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-
abén, 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 holotype from Corumbataf, 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 the locality.
Description. Female holotype. Carapace
red-brown 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
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 corumbatai_ 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).
Distribution.
zil (Map 3C).
Specimens Examined. No other specimens have
been collected.
Santa Catarina State, Bra-
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 Falc6n, 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 aah white longitudinal rectangle.
Carapace with tubercles and _ scattered
long white setae (Figs. 121, 122). Legs
aan 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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
aul wwlore, SJ waren, dette! G0), toorda 2h),
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.
IQ).
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; Map 3B
Mastophora carpogastra Mello-Leitao, 1925: 460.
Two female syntypes from Rio de Janeiro, Brazil,
in MNE 672, examined. Mello-Leitaio, 1931: 72,
na Gh, IS, De
Ghee an fagoides Vellard, 1926: 327, figs., 2,
egg sac. Female holotype from Butantan, Sao Pau-
lo, Brazil, in IBSP, lost. First synonymized with car-
pogastra by Mello-Leitaéo (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-
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
mim, 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 oilers: In
the cleared epigynum, the atria separate
ventrally (Figs. 133, 134).
The egg sac elke 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, ach 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.
Distribution. 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). Sdo Paulo: Agua da Figueira Maracai, 9 Feb.
1967, 12 (G. Brisolla, IBSP 2064); Barueri, Apr.
1963, 1°, 5 egg sacs (K. Lenko, MZSP 3069); 16 Mar.
1966, 1 imm. (K. Lenko, MZSP 5265); Diadema,
June 1986, 12 (R. Snignani, IBSP 4992); Embu,
Sept. 1982, 12, egg sac (A. L. Prestes, IBSP 3501);
Jandira, May 1980, 1? (C. Luiz, IBSP 963); Osasco,
25 Mar. 1974, 12 (F. Ramirez, IBSP 2721); Pacaem-
bu, Nov. 1942, 12 (Braudas, MZSP 357): Perdizes,
MASTOPHORA ° Levi 341
26 Aug. 1951, 12 (H. Camargo, MZSP 7496); Ribei-
rao Pires, May 1975, 12 (F. B. Lopes, IBSP 3591);
Rio Claro, July 1941, 1° (P. Pereira, MZSP 4552); 4
May 1942, 1 imm. (Clareteano, MZSP 4402); Saco-
ma, 7 Sep. 1943, 1 (J. Lima, MZSP 4403); Sao Pau-
lo, July 1921, 12 (MZSP 8068); Feb. 1928, 1° GE
Lima, MZSP 8070); 31 Jan. 1934, 12 (M. Oliveira,
IBSP 1920); 26 Bebe 1936, 12 (S. Remetente, IBSP
3598): 3 Oct. 1951, 12 (R. Vieira, IBSP 586); 17 Mar.
1955, 12, 2 egg sacs (J. Navas, IBSP 1176); June
1960, 22 (J. London, IBSP 1552); June 1960, 1° (L
Zodiygansky, IBSP 1516); 21 IDS 1960, 12 (W. Ge
drade, IBSP 1618), 10 July 1962, 12 (R. R. Guidug-
lin, IBSP 1775); 21 Dec. 1961, 12 (F. V. Boas, IBSP
1620); 18 July 1962, 12 (E. Botelhiat IBSP 1776):
Apr. 1963, 12 (Merck Co., IBSP 1834); 2 Aug. foeee
1¢@ (S. Remetente, IBSP 1949): 16 Dec. 1971, 19 a
Rafael de Simone, IBSP 309); June 1975, 12 (J. S
Gomes, IBSP 3594): Feb. 1976, 12 (M. Uchiyama,
IBSP 3597): July 1975, 12 (G. P. Treu, IBSP 3595);
Feb. 1976, 12 (E. I. Yamane, IBSP 17789): Feb.
1982, 12, egg sac (D. Zammataro, IBSP 3025); 26
eb. 1982, 1, egg sac (M. C. Franco, IBSP 14279);
2 Jan. 1986, 1° Gn Steiner, IBSP 8471); 21 Mar.
Aa 12 (D. R. Bizzachi, IBSP 14424): 1 Feb. 1991,
(J. Batista, IBSP 14398): Oct. 1992, 12 (C. M
Nerici, IBSP 5827); 7 Oct. 1996, 12 (A. Fallatti,
IBSP 14192); 11 Aug. 1997, 12 (S. M. Carnelho,
IBSP 14012); 18 Mar. 1998, 12 (A. Pastore, IBSP
16208): Brooklin, Feb. 1962, 12 (L. Travassos, MZSP
4347); Hato Museu Paulista, Feb. 1951, 12 (C. Ra-
bello, MZSP 6609); Ipiranga, Nov. 1906, Oct. 1912,
2@ (Ihering, L. M. Torre, MZSP 3047); Mar. 1924 (J.
Lima, MZSP 3048); 15 Mar. 1961, 12, 5 egg sacs
(Almeida and Cautero, MZSP 4359); 4 May 1961,
19, 2 egg sacs (N. C. Oliveira, MZSP 4340); 23 Jan.
1984, 1 imm. (C. R. F. Brandéo, MZSP 435); Magi
das Cruzes, Ranch das Carmelitas, June 1976, 12 (C.
Torrus, S. Filho, MZSP 11433). Santa Catarina: Ca-
ceador, 1982, 12 (D. Lorenzato, IBSP 3539). Rio
Grande do Sul: Ee Alegre, 12, 1 egg sac (P. Buck,
MNRJ 1831); 7 July 1986. 12 (S. Oresco, MCN
15236): S. Leopoldo, 24 May 1964, 12 (C. Valle,
MZSP 4233): 14 Oct. 1965, 12 (C. Valle, MZSP
5422).
Mastophora seminole new species
Figures 135-141; Map 2G
Holotype. Female holotype from Hollendale, Bro-
ward Co., Florida, 11 June 1987 (W. Birch), in
FSCA. 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
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
mim, 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. yeargani, 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,
2. 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, ister al
eyes on bulges. Abdomen light orange-
brown with darker area anteriorly beeecen
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,
B35) paide in thoracic region, 2.3 wide at lat-
eral eyes. First femur 3 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,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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
ae 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, 12
(AMNH). Holquin: Bafies, 1-3 Aug. 1955, egg sac (A.
F. Archer, AMNH).
Mastophora hutchinsoni Gertsch
Figures 153-168, 453, 454; Map 2E
Cyrtarachne cornigera:—McCook, 1890: 98, 99, fig.
81, egg sacs. Kaston, 1948: 231, figs. 741, 742, 2039
(misidentification).
Mastophora hutchinsoni Gertsch, 1955: 236, pl. 6, fig.
3, text figs. 10-14, 39, 47, 48, 2, 5. 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
2
cen
S
3
Figures 153-168. Mastophora hutchinsoni Gertsch. 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,
172, carapace and abdomen. 171, dorsal, with male. 172, lateral. 173-175, epigynum. 173, ventral. 174, posterior. 175, 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
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
min, 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 male 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 compar ed with
those of aliver 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-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
cidentalis L.) and wild cherry (Prunus 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 Ate 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, 1d
(E. B. Bryant, MCZ). MASSACHUSETTS 1 (San-
born, MCZ). Middlesex Co.: Pepperell, 15 July 1978,
limm. ¢, 1.5 mm (F. J. Murphy, JM). RHODE IS-
LAND Providence, 12 (N. Banks, MCZ). CON-
NECTICUT Hartford Co.: Rocky Hill, 23 Oct. 1940,
12 (A. Morgan, USNM). NEW JERSEY Burlington
Co.: Riverton, Sept. 1926, 2 imm., 1d (R. J. Sim,
OSU). Hunterdon Co.: White House Station, Sept.
1917, 12 (J. J. Brochon, USNM). Morris Co.: Dover,
8 Dec. 1950, egg sacs (USNM). OHIO Delaware Co.:
5 km W of Deen Dam, 40.37°N, 83.10°W, Oct.
2001, 22, egg sacs (R. Bradley, OSU). DISTRICT
OF COLOMBIA Washington, yard, 18 Oct. 1920, 12
(USNM). VIRGINIA Falls Church, 52, 2 imm. (N.
Banks, MCZ). Augusta Co.: Augusta, 14 June 1976,
13; 6 Oct. 1976, 12 ( (J. P. McCaffrey, CNC). SOUTH
CAROLINA Anderson Co.: Simpson Agric. Exp. Sta-
tion; LOT July 197385 di 13 eAug 19708 ho SON Seps
1978, 12 (G. Lee, CUAC). KENTUCKY Fayette Co.:
Lexington, 7 Sep. 1990, 12 (K. V. Yeargan, MCZ); 20
Sep. 1990, 19 (K. V. Yeargan, KVY); 2 Oct. 1999, 1°
(K. V. Yeargan, MCZ). Clark Co.: Feb. 1996, egg sac,
43 (K. V. Yeargan, KVY). TENNESSEE Ashburn, 30
mi. N. Nashville, 17 July 1933, 1d (W. Ivie, AMNH).
MICHIGAN Livingston Co.: E. S. George Reserve,
22 July 1951, 1 imm. (H. K. Wallace, FSCA). IN-
DIANA Putnam Co.: Greencastle, 12 (N. Banks,
MCZ). ILLINOIS Champaign Co.: Univ. Illinois ap-
ple orchard, Sept. 1993, 12 (S. D. Gaimari, INHS).
Jackson Co.: 8 km S Carbondale on avocado plant,
Oct. 1976, 12 (JAB). MINNESOTA Hennepin Co.:
Minneapolis, 1 Nov. 1931, 12 (W. J. Gertsch,
AMNH).
Mastophora cornigera (Hentz)
Figures 169-182, 455; Map 2G
Epeira cornigera Hentz, 1850: 20, pl. 3, fig. 8, &.
Immature female holotype from Alabama, de-
stroyed. Hentz, 1875: 123, pl. 14, fig. 8, @.
Cyrtarachne bicurvata Becker, 1879: 77, pl. 2, figs.
16-19, 2. Female holotype from peach tree, Don-
aldsonville, Louisiana, in IRSNB, examined. First
synonymized by Marx (1890).
Cyrtarachne cornigera:—Keyserling, 1880: 300, pl. 4,
fig 4502 MeCocle 1890: 98. fig. “80 (in part).
Ontlauntns cornigerus:—Marx, 1890: 541. McCook,
1894: 197, pl. 12, fig. I, @.
Glyptocranium cornigerum:—Simon, 1895: 882, 885.
Bonnet, 1957: 1996.
Mastophora cornigera:—Mello-Leitao, 1931: 70, figs.
9, 20. Gertsch, 1955: 233, pl. 6, fig. 2, text figs. 1-5,
37, 41, 42, 2, 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—
14.0 mm, males 1.6—1.7. The illustrations
were made from the female from Alabama
(Figs. 169-175) and males from California
MASTOPHORA ° Levi 345
(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. yeargani 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 Jat-
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°
(K. V. Yeargan, KVY). TENNESSEE Ashburn, 30 mi.
N Nashville, 17 July 1933, 12 (W. Ivie, AMNH). AL-
ABAMA Mobile Co.: Mobile, 12 (N. Banks, MCZ):;
1932, 12 (H. PB. Loding, MCZ). LOUISIANA East
Baton Rouge Par.: Baton Rouge, Apr. 1916, 12 (New-
ell, MCZ). Orleans Par: New Orleans, 1918, 12 (H
E. Hubert, USNM); 1 Oct. 1935, 12 (T. E. Snyder,
MCZ); 26 Sep. 1936, 12 (J. N. Cowanloch, USNM).
TEXAS Travis Co.: Shellberg Tract, 30°25'N,
97°52’W, 18-19 Apr. 1994, 3d (Dunlap et al.,
TAMU). Galveston Co.: Texas City, 1921?, 2° (S. W.
Bilsing, MCZ). San Patricio Co.: Welder Wildlife
Refuge, 11.8 km NE Sinton, 17 Oct. 1967, 1d (C
Parrish, CAS). Hidalgo Co.: Edinburg, Oct. 1934, 39,
1d; 1935, 12 (S. Minleie AMNH): 7 Dec. 1935, 14
(M. Welch, AMNH), 2 imm., 16 (S. Mulaik,
AMNH): 18 km SE Pharr, Santa Ana Wildlife Ref-
uge, 1 Oct. 1977, 12 (O. Ahrenholtz, AMNH); 20
346
Deer 983n ll 2a(M Stowe, MKS); 27 June 1984,
12 (M. K. Stowe, Anes ee 10 Apr. 1986,
egg sac, 70 imm., 596 (D. A. Dean, TAMU). Cam-
eron Co.: Brownsville, 9 Apr. 1986, hatched 2 June
1986, 1 egg sac, 64 imm., 62d (D. A. Dean, TAMU);
Harlingen, Feb. 1980, 12, 64 imm., 63d (C. W. Ag-
new et al., TAMU); E of Harlingen, 3 Jan. 1936, 12
(L. I. Davis, M. Stegmeier, AMNH). ARIZONA Mar-
icopa Co.: Phoenix, Apr. 1941, 12 (AMNH). CALI-
FORNIA Contra Costa Co.: Walnut Creek, Sydney
Drive, July 1992, 12, 1d (T. Trosin, J. Fraser, CAS).
Santa Clara Co.: Palo Alto, 1914, 12 (H. Heath,
MCZ). Santa Barbara Co.: Santa Barbara, 5 Oct.
1948, 12 (H. Shantz, AMNH). Los Angeles Co.:
Claremont, 12 (N. Banks, MCZ); Glendale, 1°,
many 6 (C. E. Hutchinson, AMNH, MCZ); Los An-
geles, 12 (MNHN, 3059): 10 Oct. 1942, 5d Qe EL:
Branch, AMNH); East Los Angeles, 1943, 12 (C.
Cowles, AMNH); Malibu, Nov. 1968, 12, 3 egg sacs
(USNM); Westwood Village, Aug.—Oct. 1942, 12 (P.
Verrity et al., FSCA). Orange Co.: Santa Ana, 4d (R
K. Bishop, USNM); San Juan Capistrano, 25 Sept.
1952, 1?, 7 egg sacs (R. E. Ryckman, AMNH). San
Bernardino Co.: San Bernardino, 1880, 12 (J. B. Par-
ish, MCZ). San Diego Co.: San Diego, 22 (USNM),
4 Oct. 1974, 1° (D. Bishop, USNM); San Diego,
Vista, 8 June 1989, | imm. (J. W. Schott, MCZ); Chu-
la Vista, 5 Dec. 1981, 12 (H. V. Weems, FSDA): 10
mi. NE Ramona, 22 July 1982, 19 (J. Halstead, DU);
Lakeside, 12 (C. Kingery, USNM); Feb. May 1968,
egg sacs, dd (C. Kingery, USNM). MEXICO Baja
California Sur: 44 km W La Paz, 0.2 km S km 44,
on Highway 1, 31 Dec. 1978, 12 (D. Weissman, R.
Love et al., CAS). NICARAGUA Managua: Mateare,
12 Sep. 1995, 12 (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, 2, d. 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, later al eyes on bulges. Abdomen
mn 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,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
third 2.3, fourth 3.7. Length of first patella
and tibia 1.3 times 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 from
M. phrynosoma and M. hutchinsoni by
having only a minute terminal apophysis
(Figs. 190, 191). It differs from M. phry-
nosoma 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 Baldwin 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. WM. fasciata Reimoser. 194-200, female. 194, 195, carapace and chelicerae. 194, frontal. 195, lateral. 196,
197, carapace and abdomen. 196, dorsal, with male. 197, lateral. 198-200, epigynum. 198, ventral. 199, 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.
345
Sep. 1949, 1d allotype (A. F. Archer). KANSAS
Douglas Co.: 22 Sept. 1948, 12 (R. H. Beaner,
AMNBE).
Specimens Examined. SOUTH CAROLINA
Charleston, Oct. 1941, 12 (E. B. Chamberlain,
USNM). FLORIDA Alachua Co.: Devil’s Millhopper
State Park, reared spring 1992, 4d (M. K. Stowe,
MKS). Hillsborough Co.: Thonotosassa, 11 Dec.
1969, 1? (D. A. Vaughn, FSCA). ALABAMA Mobile
Co.: Mobile, 12 Nov. 1939, 12 (A. C. Cole, AMNH);
Mt. Vernon, Oct. 1941, 1? (H. P. Léding, AMNH).
Mastophora fasciata Reimoser
Plate 1; Figures 194-204, 457; Maps
2F, 3A
Mastophora fasciata Reimoser, 1940: 356, fig. 8, &.
Female holotype, from Orosi, Proy. 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 Poz6n [Falcén], 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 abdomen and the posterior light band.
Description. Female from Costa Rica.
Carapace light orange-brown. Sternum
brown. Legs lighter brown. Abdomen
white with Tblaek 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
ILL x5) Lacon, Carapace 4.7 mm long, 5.0 wile
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
andistibial 4:6. mms) thing) 247. stountia 4.0)
Length of first patella and tibia 1.3 times
width of carapace.
Bulletin Museum of Comparative Zoology
; Vol. 157, No.5
Male 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 macroseta.
Total length 1.6 mm. Carapace 0.74 mm
long, 0.65 wide in thoracic region, 0.46
wide behind posterior lateral eyes. First
femur 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 Escazi.
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: 04) 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. Males are mature
when they leave the egg sac.
Distribution. The species is known from
Costa Rica and Venezuela (Maps 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, mesal. 214, ventral. 215, ectal.
MASTOPHORA ® Levi 349
pickeli
Wye jy?
Maa ig
i) CAIpr Ay
yr, ) Uf) WE?
We PG,
d i yy 04
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
Specimens Examined. COSTA RICA San José: San
José, 12 (Tristan, MCZ); San Antonio de Escazt,
1,350 m, May 1980, 12 (W. Eberhard $31, MCZ): 7
Sept. 1980, imm., 2, many 6 (W. Eberhard SJ1—26,
MCZ): 11 Oct. 1980, 12 (W. Eberhard $31, MCZ).
Puntarenas: Parrita, 30 m, 12 Jan. 1987, 32, 2d (W.
Eberhard FN9—40ff, MCZ).
Mastophora dizzydeani Eberhard
Figures 205-215; Map 3A
Mastophora dizzydeani Eberhard, 1981: 144, figs.
1-9, 2, d. 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—
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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.
PAUL).
Males have a smaller median apophysis
(Fig. 213) than that of M. fasciata, in ectal
view surrounding a rounded space (Fig.
PAS)
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 frugi-
perda (a sugar cane pest) and Leucania sp.
(Eberhard, 1977, 1981).
Distribution. Colombia to northern
Peru (Map 3A).
Specimens Examined. COLOMBIA Valle: Sof
Cali, on plants, 6 June 1948, 12 (E. M. Poulsen,
ZMUC); nr. Cali, Jan. 1977, imm. 2 (W. Eberhard,
MCZ); Aug. 1977, 12, 4d (W. Eberhard, MCZ);
Lago Calima, 1,400 m, 19 Nov. 1977, 12 (W. Eber-
hard EG 3-20, MCZ); Rio Tulua, 1,100 m, Aug.
1977, 26 (W. Eberhard, MCZ). PERU Piura: Mal-
lagra, Rio Chira, 8 June 1941, 22, 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, 2. Female holotype from Tapera, Pernam-
buco, Brazil, in MNRJ, 395, examined. The specific
name is a noun in apposition after the locality.
Roewer, 1942: 901. Platnick, 2001.
Glyptocranium 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
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 similar 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, pl. 1, @.
Female holotype from Tapera, Est. Pernambuco,
Brazil, in MNRJ no. 00394, examined. Mello-Lei-
tao, 1931: 72, figs. 2, 14, 9. Roewer, 1942: 955.
Platnick, 2001.
Glyprocranium 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-
MASTOPHORA ° Levi 351
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.
929).
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-Leitéo, 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,
fingerlike (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
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 stawles light brown, darker ventrally,
ith white Stace 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 Te 3mm, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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; Map 3D
Holotype. Female yolotype from Emas, Pirassunga,
Est. Sao Paulo, Brazil, 2 Nov. 1952 (Pietracatelli,
Werner, and Dionisio), in MZSP, 4339. The specific
name is an arbitrary combination of letters.
Description. Female holotype. Carapace
dark brown. Chelicerae light brown. La-
bium, endites, sternum light brown. Coxae
and distal leg articles began with indistinct
darker Bigtches! 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. Mastophora 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. MM. 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
a0
CORY
2,
S
longiceps
Figures 244-250. OM. 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. ypiranga 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
legs, and anterior of abdomen; the abdo-
men is all dark, perhaps because of poor
preservation. The illustrations were from
the holotype.
Diagnosis. Mastophora piras is distin-
guished from M. melloleitaoi 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: Gavernador Va-
ladares, 14/15 Oct. 1981, 1 imm. (N. Sorkin, T. Spitz-
man, AMNH). Sdo Paulo: Sio Bernardo, Nov. 1926,
12 (H. Bakkenist, MZSP 8069).
Mastophora ypiranga new species
Figures 251-257; Map 3E
Holotype. Female holotype from Ypiranga, Cap. [city
of Sido 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 brown.
Coxae and distal leg articles light brown.
Abdomen anterior black, posterior white
(Figs. 253, 254); venter grayish white with
white square. Carapace with low tubercles,
with some short white setae on sides (F igs.
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
mim, 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—
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
12.7 mm. The illustrations were made
from 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,
12 (E. J. Hambleton, AMNH). Sdo Paulo: Sao Paulo,
July 1928, 12 (R. Cassalo, IBSP 4530). Santa Catar-
ina: June 1919, 12 (Luederwaldt, MZSP 8067).
Mastophora yacare new species
Figures 258-264; Map 3E
Holotype. Female holotype from Rincori del Yacaré,
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. Abdomen 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, posterior.
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
eet fis yi?
fae % 258 ae % 259 Ps
abalosi
ae yacare
260
as
n n
n
a
(498 Phe
& aoe
(ger eo 4
5 \
Ay ie
- v 28I 282 oe
Neier eth 2s 182
ON
extraordinaria
278 283
285
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
2.0, fourth 3.3. Length of first patella and
tibia 1.3 times width of carapace.
Males are not known.
Diagnosis. Mastophora yacare is distin-
guished from M. melloleitaoi by its epigyn-
um 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, 2. 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
Badez, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Mastophora comma Baez and Urtubey
Figures 269-273, 459; Map 3E
Mastophora comma Baez and Urtubey, 1985: 3, figs.
1-9, 2. 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 structure 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-Leitdoi Canals, 1931: 20, figs. 1-4,
pl. 2, fig. 4, 2. Female holotype, from Rosas, Prov.
Buenos Aires, Argentina, in MACN, examined.
Mastophora_ mello-leitéo:—Mello-Leitao, 1931, 73:
Ine Oy ILIA Se
Mastophora mello-leitaoiae:—Roewer, 1942: 900.
Glyptocranium melloleitaoi:—Bonnet, 1957: 1998.
Mastophora melloleitaoi:—Platnick, 2001.
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
andatibial4:oyamm) thirds 2. 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, 12
(B. P. Mann, MCZ). Paranda: Curitiba, 21 Jan. 1965,
1 imm. (C. Valle, MZSP 4326). Rio Grande do Sul:
Rodeio Bonito, Bagé, 10 Feb. 1967, 12 (C. de Oliv-
eira, MCN 473); Santa Maria, 9 May 1973, 12 (D.
Link, MCN 01659). ARGENTINA Cordoba: Cruz
Alta, 15 Aug. 1948, 12 (J. P. Duret, MACN). Buenos
Aires: Ascension, 12 (B. Gerschman, MACN): Bal-
carce, 16 Feb. 1950, 12 (Cuccioli, MACN):; Burzaco,
MASTOPHORA ® Levi 357
12 (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 Holmberg, 1876: 113. Fe-
male from Buenos Aires, Argentina, lost. Bréthres,
1909: 163, figs. 1, 2, 2, egg sacs. Canals, 1931: 17,
figs. 1-5, pl. 1, fig. 1. Mello-Leitao, 1931: 70, figs.
7, 19. Roewer, 1942: 900. Platnick, 2001.
Glyptocranium extraordinaria:—Bonnet, 1957: 1997.
?Mastophora cinerea Mello-Leitaéo, 1943: 105, fig. 4,
imm. Immature holotype from Cordoba, Argenti-
na, in MLP, examined. DOUBTFUL NEW SYN-
ONYMY.
Mastophora intermedia Mello-Leitao, 1945: 240, figs.
14-17, 2. One female holotype from Pindapoy, Mi-
siones Prov., Argentina, in MLP, examined. Brig-
noli, 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. Female from Chascomis.
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
mm, patella and tibia 5.5, metatarsus 3.5,
tarsus 1.2. Second patella and tibia 4.5
mim, third 2.7, fourth 4.1. Length of first
patella and tibia 1.1 times width of cara-
pace.
Males are not known.
358
Variation. The epigynum is variable in
width and the abdominal humps may be
wider or higher. Total length of females
95-14 mm. The illustrations were made
from the female from Chascomis.
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 epigynum is flat dorsally, sclerotized,
and darker than ventrally (above on Fig.
286); it is raised in the middle in M. mel-
loleitaoi. The atria approach each other
(Fig. 287), unlike those of M. melloleitaoi
(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
Province, west to Chaco and Cérdoba
provinces, Argentina (Map 3F).
Specimens Examined. BRAZIL Rio Grande de Sul:
Canela, 12 Feb. 1966, 12 (A. Lise, MCN 0119): Gar-
ibaldi, 30 Oct. 1974, 12 (O. ame MCN
2381). URUGUAY Montevideo, 1? (J. Canosa,
MACN 4182); 12 (E. Cordero, MNRJ 14011); Pun-
tas Arroyo Laureles, Tacuaremb6, 1 imm. (FCMU
293): Treinta y Tres, 20 Aug. 1971, 1 2 (FCMU 295).
ARGENTINA Chaco: Saenz Peja, Sept. 1933, 19
(B. Ohneiser, MACN 31331). Cordoba: Cérdoba, 22
M. J. Viana, AMNH, MACN 1106); Calamuchita,
Dec. 1940, 49 (J. M. Viana, MACN 1005): Dec.
1941, 22 (J. M. Viana, MACN 1106); Agus do Oro,
Mar. 1940, 1 imm. (J. A. De Carlo). Buenos Aires:
Buenos Aires, 12 (Scly., MNHN 23388); Chascomits,
Oct. 1934, 12 (I. Dor, MACN 35983); Cap. Federal,
limm. (E. Pizarro, MACN 12782): Florencio Varela,
1 May 1949, 1° (O. de Ferrarini, MLP 13549); Gon-
zales Catan, 6 June 1949, 12 (Touson); Hurlingham,
Jan. 1954, 12 (Giai, MACN 4359); La Plata, 12 (M.
Birabén, MLP 16178); 12 (M. Birabén, BMNH):
Moreno, Feb. 1939, 12 (Schiapelli, Gerschman,
MACN); 14 Nov. 1943, 12 (S. M. Doello Jurado;
MACN 1361); Rosas, 1930, 1° (J. B. Daguerre,
MACN 4185).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Mastophora leucabulba (Gertsch),
new combination
Figures 288-295; Map 2F
Agatostichus leucabulba Gertsch, 1955: 250: figs. 34,
38, 40, 2. Female holotype from Harlingen, Texas,
in AMNH, destroyed.
Agathostichus leucabulba:—Brignoli, 1983: 255.
Agathostichus leucabulbus: pisincle 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 Ibvaeseornfislh at base. Labium, en-
dites yellowish. Sternum yellowish. Coxae
yellowish and distal leg articles yellowish
to light brown with fait brown markings
on fins 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 iedian bulb flanked by a pair of
sniel bulbs on the side, behind bifid
horns, sides with dark warts and cephalic
area coarsely roughened, hidden by mat of
wooly hairs. betel 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
mim long, 2.9 dee 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 times 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
leucabulba
Figures 288-295. Mastophora leucabulba Gertsch. 288-291, female, after Gertsch. 288, carapace and chelicerae, 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. WM. alvareztoroi lbarra and Jiménez 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 right palpus. 305, dorsal. 306-307, left
palpus. 306, mesal. 307, ventral.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
360
ego, 29 Apr. 1895, 2d, imm. doubtful determination
(USNM). Wilson Co.: Floresville, 29 Apr. 1895, 1
imm., 4d, 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 Jiménez new species
Plate 1; Figures 296-307; Map 2F
Holotype. Female holotype from Rancho Alejandria,
Municipio Estaci6n 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: 173, fig.
ES SR photos
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 aise leg articles
yellowish; first femur with fine black ring
and distally with ventral dark patch. Nie
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. Chelicerae
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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
mim, 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
shout half length of median horns, sepa-
ration beueen 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 ne 1.1 mm, pa-
tella and aie 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. WM. 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
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. leucabulba 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), Rind 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 | 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 Estacién Juarez, 25 Sept. 1975, 12 (M.
Alvarez del Toro, MLJ); Finca Inlanda, 870 m,
15°10'N, 92°21'W, 65 km NNW of Tapachula, 12
Aug. 1987, 1d (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, 12 (A. Ventura, ECOTAR).
Specimens Examined. TEXAS Hidalgo Co.: Santa
Ana Natl. Wildlife Refuge, 18 Dec. 1983, 12 (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).
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 the
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) 422 imi) third)! 3a tomntamors
Length of first patella and ie 1.7 times
width of carapace.
Males are not known.
Diagnosis. This species is distinguished
from M. alvareztoroi by the longer cara-
pace tubercles and by having Ths 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,
Bahia, Brazil, in MNHN, 8486, examined. Mello-
Leitao, 1931: 67. Gertsch, 1955: 250.
Agathostichus leuwcacantha:—Simon, 1897: 473.
sRoewer 1942: 900.
Agathostichus leucacanthus:—Bonnet, 1955: 182.
Platnick, 2001.
Note. In his publications, Simon (1895,
1897) did not include the female symbol
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 valid 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, wench lateral pair on a
swelling. Median Benue 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
MASTOPHORA ® Levi 363
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. Dinner ror roi by having the ab-
domen rounded behind, sphereas 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 Orgaios, Organ Moun-
tains |, 1902. 19 (iss 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
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 brown with dark
streaks and speckles, and many downy
white setae (Figs. 322, 323). Chelicerae
patchy brown. Abyann: endites, sternum
dark brown. Coxae light brown and distal
leg articles yellowish iin temo brown
rings. Abdomen light brown, 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 ede
in thoracic region, 1.8 wide Benne 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
Dy rowiada Sex
Males are not known.
364
Diagnosis. Mastophora brescoviti 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 Holmberg, 1876: 143. Female
from Boradero [Prov. Buenos Aires], Argentina,
lost.
Mastophora conifera:—Canals, 1931: 18, figs. 1—5, pl.
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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
(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 Fé and Buenos Ai-
res provinces, Argentina (Map 3G).
Specimens Examined. ARGENTINA Santa Fé: Co-
lonia Macias, Nov. 1942, imm. shriveled (J. M. Viana,
MACN). Buenos Aires: Tigre, 1902, 12, once dried
up (J. Brethes, MACN 5896).
Mastophora corpulenta (Banks)
Figures 336-342, 461; Maps 2F, 4A
Ordgarius corpulentus Banks, 1898: 251, pl. 15, fig.
8. Female holotype from San José 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, 2. Female holotype from Zamorano [?Zambra-
no], Honduras, in AMNH, examined. Brignoli,
1983: 274. Platnick, 2001. NEW SYNONYMY.
Glyprocranium 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
G. Marx before being turned over to
Banks after Marx’s death, and Marx 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. MM. 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
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
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.
Diagnosis. 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
AA).
Specimens Examined. NICARAGUA Leé6n, Aban-
gasca, 13 Dec. 1994, 1 subadult 2 (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. Secu 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
light tips (Figs. 343, 344). Median eyes on
bulge, lateral eyes on bulges. Abdomen
slightly wider than long wath small humps
(Figs. 345, 346). Total length 7.7 mm. Car-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 History. From Y. Lubin (per-
sonal correspondence): “#319. nocturnal
araneid on orb web. During day sits on
twigs. 1 female, 1 juvenile. [NB: maybe 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 Rell outwards, flexed. No Taelew:
Spider Smerchcs legs forward when I hum,
then moves to edge of leaf and adopts
same posture.”
Distribution.
4A),
Galapagos Islands (Map
Paratypes. Galapagos Islands: Isabella Island, Ta-
gus Cove, on Croton scouleri at night, 13 May 1983,
1 imm. (Y. Lubin 510, MCZ).
Mastophora escomeli new species
Figures 351-361; Map 4A
Escomel, 1918,
Glyptocranium gasteracanthoides:
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 the 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. WM. 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. MM. 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
Figures 380-386. MM. 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
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 curvature. 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. a mm, patella and tibia Wat
metatarsus 6.3, tarsus 1.6. Second patella
ands tibia 4.8 mm: third: 227. 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 a 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 free patella and tibia
15> e/atimmes Sadeh of carapace. The fe-
male from Ica is more setose, with long
setae on the legs and many shorter setae
on the Aileen: 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 escomeli is dis-
tinguished from M. gasteracanthoides by
having a swelling betwecn the slits on the
posterior of the “epigynum (Fig. 356) and
having the atrium ventral to the seminal
receptacles (Fig. 357). In M. gasteracan-
Bulletin Museum of Comparative Zoology
, Vols ISG, INO. 5
thoides, the slits are in a shared depression
and the atria are dorsal to the seminal re-
ceptacles (Figs. 415, 416).
Natural History. 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, 12 (N. Llerana Martinez, MUSM). Arequipa:
Arequipa, 1912, 22 (E. Escobal, MNHN); 12 (E.
Escobal, MACN 4198).
Specimens Examined. PERU Ica: Ica, 1992, 1°
(Cascavilea-Rubio, MACN). La Libertad: Cerro
Campana, N Trujillo, 23 May 1989, many 6, 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 obtusum:—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-
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, Broa, in MZAQ
no. AO105, examined. Brignoli, 1983: 273. Platnick,
2001.
Note. The holotype was embedded in
difficult-to-remove fungal mycelium.
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, whites brown. Sternum brown.
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 having long wide
humps and carapace with fabercles on
sides and by having the atria of the epi-
gynum ventral to fie 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.
MASTOPHORA °® Levi 369
Distribution. Rio de Janeiro and Sao
Paulo states, Brazil (Map 4C).
Paratypes. BRAZIL Rio de Janeiro: Santo Anténio,
Rio Bonito [22°42'S, 42°37'W], 1933, 12 (S. Reme-
tente, IBSP 418). Sdo Paulo: ?Campinas, July 1982,
12 (C. Froelich, IBSP 4968).
Mastophora holmbergi Canals
Figures 373-379; Map 4C
Mastophora Holmbergi Canals, 1931: 22, figs. 1-5, pl.
3, fig. 5; pl. 4, figs. 7, 8, 2. Female from km 701,
Santiago del Estero, in MACN, 24133 [7140], ex-
amined.
Mastophora holmbergi:—Mello-Leitao, 1931, 73, figs.
10, 21, 2. Roewer, 1942, 900. Platnick, 2001.
Glyptocranium 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 Pant 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 made in thoracic region, 3.0 wide
at later al 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—
Se) mana, ene ‘ilmecaciens were made
from the female holotype.
Diagnosis. Unlike Mastophora reimo-
seri, ih holmbergi has long first legs. In
the epigynum fhe atria are sable 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, 12 (Cranwell-Giai, MACN
1630).
370
Mastophora reimoseri new species
Figures 380-386; Map 4C
Holotype. 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 erennict humps and short se-
tae, base of humps 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
mim, 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 ies musical note—shaped Pree of the
slits in the epigynum (Fig. 385), and by
having shorter legs than M. holmbergi.
Distr ibution. Known only from Roun:
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, pl. 3,
fig. 6, 2. Female holotype from La Rioja, Argen-
tina, in MACN, 5260, examined. Mello-Leitio,
1931: 73, figs. 11, 22, 2. Roewer, 1942: 901. Plat-
nick, 2001.
Glyptocranium satan:—Bonnet, 1957: 1998.
Description. Female from Cérdoba.
Carapace dark brown with many short
light setae and narrow white rim. 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
eyes on bulge (Figs. 387, 388). Abdomen
humps very 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. Seas legs beige. Abdomen
whitish. Carapace with four tubercles: ab-
domen with two humps. Total length ed
mm. Carapace 0.92 mm long, 0.81 wide in
thoracic region, 0.58 wide a lateral eyes.
First fonatin 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 times cara-
pace width in females 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 Cérdoba Province, Argentina, a
female from La Rioja Province (Fig. 394),
a female from Tucumaén Province (Fig.
395), and of the male from Rio Grande do
Sul, Brazil.
Diagnosis. Mastophora satan is separat-
ed fon M. gasteracanthoides and M. dia-
blo by the long first tibia and metatarsus,
each 9.7 mm 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-
fhoiles but differs in ventral view, where
MASTOPHORA ° Levi 371
ae M10) ds
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
M. gasteracanthoides has a distinct sclerite
a short distance anterior to the posterior
edge (Fig. 414), whereas M. satan has the
whole median area slightly sclerotized
(Fig. 391).
The median apophysis of the palpus of
the male (Fig. 398) is more curved 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, 12 (B. Pickel, MNRJ 391). Bahia: Feiora de San-
tana, July 1994, 12 (S. D. Cunha, IBSP 16246). Sao
Paulo: Santo André, 16 June 1965, 12 (L. Daga,
IBSP 1931); Seminario Santa Terezinha, Tieté, 5 May
1953, 12 (IBSP 887); Sao José do Rio Preto, 9 Feb.
1964, 12 (Vizotto, MZSP 3471): 1 June 1964, 1 egg
sac, 12 (Vizotto, MZSP 3470). Santa Catarina: Bla
menau, 12 (NHMW). Rio Grande do Sul: 12 (P.
Buck, MNRJ 41644); Rodeio Bonito, Bogé, 5 June
1980, 12 (E. W. Aguiar, MCN 9103); Canela, 10 Feb.
1966, 29 (A. A. Lise, MCN 0752); Sao Leopoldo, 5
Mar. 1965, 12 (C. Valle, MZSP 4797); Porto Alegre,
12 Apr. 1926, 12 (R. Gliesch, MNRJ 392); Parque
Zoologico, Sapucaia do Sul, 9 Dec. 1985, 22 (A. E.
Tovares, MZSP 14079); Porto Alegre, 17 Mar. 1955,
1 imm. (T. de Lema, MCN 01628); 16 June 1963, 1°
(A. Lise, MCN 01820): 6 Oct. 1988, 12 (R. Villanova,
MCP 105); Belem Velho, Porto Alegre, 17 July 1979,
12 (V. Mott, MCN 2608): Morro Santana, Porto Ale-
gre, 5 May 1984, 1d (S. M. Silva, MCN 12204); 13
Sept. 1984, 12 (A. A. Lise, MCN 29426); Santa Ma-
ria, Aug. 1986, 12 (MCP 10340); Santo Anténio da
Patulha, 30 Oct. 1980, 12 (T. K. Moreira, MCN
9456). URUGUAY Villasboas, 1953, 12 (L. Lecour,
FCMU); Paso del Cerro (Artigas), May 1956, 192 (C.
Fuques, FCMU)); Artigas, Sept. 1959, 12 (C. Fuques,
FCMU): Mar. 1965, 12 (C. Fuques, FCMU 1965);
Ruta 3, Salto, 3 Aug. 2001, 1 (V. Vazquez, Williams,
FCMU 562); Cuareim, Espinillares, Artigas, 12 Mar.
1956, 12 (C. Fuques, FCMU 296). ARGENTINA
Misiones: 1940, 12 (Exp. Zotla. Armanini, MACN
2050). Catamarca: Sijan, Nov. 1964, 12 (Ahumada,
MACN). La Rioja: 22 (Sr. Giacomelli, MACN 4186).
Tucumdn: Tucuman, 12 Dec. 1984, 12 (FMLT 2159).
Santiago del Estero: Santiago del Estero, 20 Apr.
1958, 19 ( (J. W. Abalos, MACN); La Banda, 1958, 12
(J. Abalos, MACN); July 1958, 12 (D. Bravo,
MACN): Tabla Redonda Depto. Banda, 23 Dec.
1959, 12, egg sacs, imm. (J. W. Abalos, MACN).
Mendoza: Mendoza, 1907, 12 (E. Reimoser,
NHMW); San Rafael, Feb. 1940, 12 (D. Pereyra,
MACN 1799). Cordoba: Mina Clavero, April 1973,
12 (Stirbel, MACN). Entre Rios: Concordia, 1931,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
12 (MNBJ 57953). Santa Fé: Santa Fé, 1931, 12 (M
Birabén, MNRJ 522). Buenos Aires: Ireneo Portela,
30 April 1922, 12 (Scheimer, MACN). La Pampa:
General Pico, Feb. 1952, 12 (Williamson, MACN);
30 Mar. 1958, 22, doubtful determination (William-
son, MACN).
Mastophora diablo new species
Plate 1; Figures 399-407, 464; Map 4D
Holotype.
Chaco, Argentina,
MACN, no. 5432.
Mastophora gasteracanthoides:—Mello-Leitao, 1931:
69, in part. Canals, 1931: 19 (misidentified gaster-
acanthoides Nicolet). :
Female holotype from Colonia Benitez,
Sept. 1959 (Bachmann) in
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 Sade in thoracic region, 3.2 wide
at Teter al 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 Abie 1.5 times
wath of carapace.
Males are not known.
Variation. Total length of females 10.3—
16.7 mm. Length of feet patella and tibia
LSS etimes sate of carapace. Figures
399-405 were made from the fannie 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. 404407).
The egg sac lacks flaps and has a heavier
stalk (Fig. 464) than that of M. satan.
Natural History. A large syrphid fly (14
mm total length) was collected in Moreno,
Buenos Aires, with one adult.
MASTOPHORA ° Levi 373
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, with male. 411, 413, lateral. 410, 411, (Santiago). 412,
413, (Chillan). 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.
(ey)
~
Distribution. Northern and central Ar-
gentina to La Pampa Province (Map 4D).
Paratypes. ARGENTINA Formosa: Las Lomitas,
Oct. 1966, 12 (A. Vogt, MACN). La Rioja: 29 (Prof.
Gémez, MACN 4187). Tucumdn: El Timbé, 27 May
1952, 22 (J. Campos, FMLT 994); Los Bosques, 12
BMNH); Dept. Burruyaco, E] Haranjo, 13 June
1964, 12 (M. Inés Cortez, FML 1693). Santiago del
Estero: El Zanj6n, 5 June 1960, 12 (J. W. Abalos,
MACN); Fernandez, 4 Apr. 1960, 12 (J. W. Abalos,
MACN); Frias, 2 Oct. 1970, 12 (J. W. Abalos,
MACN); La Banda, 24 May 1958, 1¢ (J. Areas,
MACN); Los Juries, March 1959, 12 (L. Remedi,
MACN); Santiago del Estero, 18 Sept. 1958, 1 (D.
Luna, MACN); 1959, 1° (J. W. Abalos, MACN). Cor-
rientes: Bella Vista, Nov. 1944, 12 (Silberman,
MACN). Entre Rios: Basavilbaso, 12 (U. Podesta,
MACN 4189). Santa Fé: Roldan, June 1943, 12 (Es-
cuela 230, MACN). Cordoba: Argiiello, Feb. 1946,
12 (J. A. De Carlo, MACN 1657); Bajo Grande, 12
MLP 15563). Buenos Aires: Castela, 27 July 1958,
12 (Ing. Favret); Moreno, Jan. 1946, 192 (R. D.
Schiapelli, MACN 1658). La Pampa: Mira Pampa,
April 1949, 12 (C. Vigliorcho, MLP 16638); General
Pico, March 1951, 12 (C. Ballani, MLP 13642).
Mastophora gasteracanthoides (Nicolet)
Figures 7, 408-421, 464; Map 4B
Epeira gasteracanthoides Nicolet, 1849: 485, pl. 5,
fig. 7a, b, 2. Specimens from gardens and fields of
ental provinces, Santiago, Chile, in MNHN, lost.
Glyptocranium gasteracanthoides:—Simon, 1895:
882, fig. 946, 2. 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 Pefiuelas, 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 ie specimens
illustrated by Nicolet (1849) and Simon
(1895), which have higher humps. Older
specimens kept in MNHN from Chillan
(Figs. 412, 413) had longer humps but
4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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 eden 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 Chillan.
Diagnosis. Mastophora gasteracanthoi-
des differs from M. diablo 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
5
422 ie «
S Wrnagaltiels Cladomelea
akermani
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
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
oo
has a thin stalk.
Distribution. Central Chile (Map 4B).
Specimens Examined. CHILE 12 (MNHN, 114).
Coquimbo: 6 km S of La Serena, 23-30 Nov. 1961,
12, egg sac with d and imm. 2 (A. F. Archer,
AMNH#). Valparaiso: Quilput [Quilpué], 1904, 12 (C
Porter, MNHN 23457). Metropolitana: Santiago, 4
June 1947, 1 imm. (R. Donoso, AMNH); Feb. 1955,
1? (I. Pedag., AMNH); Quilicura, Oct. 1979, 12 (L.
Pena, AMNH), Aug.—Sept., 2 imm. (L. E. Pefia,
AMNH); Renca, W of Santiago, 2 Oct. 1984, 1? (L.
E. Pea, AMNH); Los Espejo, 6 Nov. 1973, 12 (C.
L. Cartagena, MCZ). Nuble: Chillan, 12 (Delfin,
MNHN, 23520).
Ordgarius Keyserling
Ordgarius Keyserling, 1886: 114. Type species by
monotypy, M. monstrosus Keyserling, 1886, from
Queensland, Australia. Neave, 1940: 453. Roewer,
1942: 902. Bonnet, 1958: 3200. Davies, 1988: 318,
mea, BO, OU 2, Cre
Euglyptila 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 Euglyptila were
described by Simon, both from immatures.
Euglyptila acanthonotata is in MNHN,
lost; the other, E. nigrithorax, 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Ordgarius magnificus (Rainbow)
Figures 422-433, 466
Dicrostichus magnificus Rainbow, 1897: 523, pl. 17,
fig. 8, 2. Holotype from Mount Kembla, New
South Wales, Australia, not examined. Roewer,
1942: 900.
Ordgarius magnificus:—Davies, 1988: 316.
Description. Female. Carapace yellow-
ish, dark brown 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, enter 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
sae 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, (Kentucky). 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 SU
yeargani
stowei
timuqua
453
hutchinsoni
archeri
corpulenta
abalosi
Ordgarius
\ 466
magnificus | '
467
Cladomelea akermani
gasteracanthoides
saian
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
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 of Mastophora (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. magnificus is spindle-shaped
(Fig. 466), that of O. monstrosus 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 Cyrtarachne longi-
pes O. P.-Cambridge, 1877: 559, from West Africa.
Neave, 1939a: 750. Roewer, 1942: 900. Bonnet,
1956: 1097.
Note. Cladomelea longipes is very simi-
lar to C. akermani.
Cladomelea akermani Hewitt
Figures 434-444, 468
Cladomelea akermani Hewitt, 1923: 63, figs. 4,5, 2.
Female holotype from Pietermaritzburg, Natal, not
examined. Roewer, 1942: 500. Leroy, Jocqué, and
Leroy, ICES I, S. 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
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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) 4976
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.
449-444), |
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). |
|
|
|
I
i
Natural History. The spider is found in
grasslands on grass of the Kwa—Zulu—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.
LITERATURE CITED
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ARCHER, A. F. 1963. Catalogo de las arafias Chilenas
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. 1910. Catalogue of Nearctic spiders. Bulletin
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BONNET, P. 1955. Bibliographia Araneorum. Tou-
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BRIGNOLI, P. 1983. A Catalogue of the Araneae De-
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MASTOPHORA ® Levi 379
CAMBRIDGE, F. P.-. 1904. Arachnida, Araneidea and
Opiliones. 2: 465-545. In Biologia Centrali-
Americana, Zoologia, London.
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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°
bisaccatum, 330
brescoviti, 361*, 363
carpogaster, 340
carpogastera, 340
carpogastra, 327*, 340, 377*
carpogastrum, 340
catarina, 335*, 339
Celaenia, 315-317
Cladomelea, 314, 316-18, 378
Coelossia, 314
comma, 355*, 356, 377*
conifera, 361*, 364
coniferus, 364
cornigera, 343*, 344, 377*
cornigera, 342
cornigerum, 345
cornigerus, 345
corpulenta, 364, 365*, 377*
corpulentum, 364
corpulentus, 364
corumbatai, 335*, 339
cranion, 349*, 351
Cyrtarachne, 315-317
Cyrtophora, 315
diablo, 32:7) Siles, SOS ont
Dicrostichus, 314
dizzydeani, 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
haywardi, 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
longiceps, 351, 353*
magnificus, 375*, 376,
Mastophora, 315-317
melloleitaoi, 355*, 356
Metepeira, 315
monstrosus, 377*
multilineata, 330
obesus, 330
obtusa, 367*, 368 |
obtusum, 368 |
occidentalis, 348 |
Ocrepeira, 315
Ordgarius, 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
>
ne
ri
A MONOGRAPHIC REVISION OF THE
ANT GENUS PRISTOMYRMEX
a . (HYMENOPTERA: FORMICIDAE)
MINSHENG WANG
97 2003
VOLUME 157, NUMBER 6
10 April 2003
(US ISSN 0027-4100)
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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
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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.)
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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
No Strac tate ek eee au eee as oe Lae RE RS 383
Tiniirahinetivor 383
(GOW ESCH OTS) meee ie ene ee A 385
Measurements and Indices _...- 385
A Brief History of the Genus Pristomyrmex _. 386
Genus eristomunmnes, Navi een ere 387
List of Pristomyrmex Names with Synonymies 392
Key to the World Species of Pristomyrmex
(Workers) eae eet et ee ee 393
Mherpunctatuss Groupee 404
Pristomyrmex OIIBU SUIS Sho hg ere 404
Pristomyrmex fossulatus (Forel)... 406
ristOmummer PUlCheTe Spos M.)) sees teres 408
Pristomyrmex punctatus (F. Smith) 410
BRiSCOMUTMEX MUA USS. (IN esate tee teseen 415
Whererbranius Group oe ee 417
Pristomyrmex cribrarius Arnold 418
itherquadridens Group) == ee 420
Pristomyrmex africanus Karavaiev 423
Pristomyrmex bicolor Emery stat. n. 425
Pristomyrmex brevispinosus Emery 428
Pristomyrmex collinus sp. n. 432
Pristomyrmex costatus sp. 0 434
Pristomyrmex curvulus SPDEAli ghosts seelel en aa 437
nristomunmex edad Morel) mars: sen 440
Pristomyrmex erythropygus Taylor 44]
PiPSHOUMUIT UD PE JUGS Os WN ee 443
Pristomyrmex foveolatus Taylor 446
Pristomyrmex UT SUTUS ES |) 1) ae eae 449
Pristomyrmex lometsmimitsssp sails eens 450
Hei StOMY TEL IMO CES CUS) S|) IN. yeeen eae eee 452
Pristomyrmex nitidissimus Donisthorpe —. 453
Pristomyrmex OCCULEUSTC Sera as sere eneaae 455
Pristomyrmex orbiceps (Santschi) 456
Pristomyrmex quadridens Emery — 459
Pristomyrmex quadridentatus (André) 463
Pristomyrmex quindentatus sp. n. 467
Pristomyrmex sulcatus Emery stat. n. _......... 469
Pristomyrmex thoracicus Taylor. 473
Pristomyrmex trachylissus (F. Smith) 474
istomu mime, (hOZOTm BOOM = 2a ieee ee 476
‘Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts 02138.
Bull. Mus. Comp. Zool.,
Pristomyrmex wheeleri Taylor 478
Pristomyrmex wilsoni Taylor 481
Mhextnispinosuss Group pe ere ee 483
Pristomyrmex bispinosus (Donisthorpe) este
Pristomyrmex browni sp. n
Pristomyrmex trispinosus (Donisthorpe) RACE 488
Mhevepisatusn Group) semis enema ene 489
Rristomueynnnen (ACCT OSUS)S)) 410.) eee ee 491
Rristomuy nme: DOLLOTL ES) \sgesee seems 492
Pristomyrmex coggii Emery —--------- 493
Pristomyrmex inermis sp. Mi. 2. 496
RristomumimexciaT SUS espe ah) ae eee 497
Pristomyrmex levigatus Emery 499
RiistomunmexiOMGtssy Sail. eeeet sesame cee 502
Pristomurenne xn ts Cid us Ist © iy aeseee amen oee eee 503
Pristomyrmex mandibularis Mann — 505
Pristomyrmex minusculus sp. M. —------ 507
Pristomyrmex obesus, Mann 2 = 509
Pristomyrmex simplex CO theae eee eer 512
DVS) 7D iRROy NAGHVIS! (Gako\ONS) 2 ee 514
Pristomyrmex profundus sp. n. -.......------- 515
ihenvmbripenniss Group yess = seen 516
Pristomyrmex fuscipennis (F. Smith) 517
PristOmmuymimen fViClett iI Iiy eee eens ee 518
Pristomyrmex pollux Domisthorjem=s =e 521
Pristomyrmex reticulatus Donisthorpe 524
Pristomyrmex umbripennis (F. Smith) 525
INiomnem INfuvdlummn 528
Pristomyrmex parvispina Emery 528
Ackaitowvled came mts eee seen eee 539
IRE LETETN CE Sie ee ts inna clean ESE ON eh ONS Rese 539
ABSTRACT. The ant genus Pristomyrmex 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
Pristom yrmex, an ant genus of moderate
size, contains 52 living species, but fossils
157(6): 383-542, April, 2003 383
384
have not been discovered. Pristomyrmex
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 I[s-
land. Lastly, one species, Pristomyrmex
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.
Pristomyrmex 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 Perissomyrmex. 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 exhibits several unusual biological
and evolutionary phenomena. The absence
of morphologically 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 morphologically
identical workers (Itow et al., 1984; Mu-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
zutani, 1980, 1982; Peeters, 1993; Tsuji,
1988a,b,c, 1990a,b, 1994, 1995; Tsuji and
It6, 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
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 specimen, 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 most
ant genera are very poorly characterized
and thus cannot be curated properly in
museum collections. Finally, I feel that
these males contain some clues for the
study of the phylogenetic relationships of
the genus.
COLLECTIONS
AMNH American Museum of Natural
History, New York, N.Y., U.S.A.
Australian National Insect Col-
lection, Canberra City, Australia
Bishop Museum, The State Mu-
seum of Natural and Cultural
History, Honolulu, Hawaii,
U.S.A.
Natural History Museum, Lon-
don, U.K.
California Academy of Sciences,
San Francisco, California, U.S.A.
Institute of Zoology, Academy of
Sinica, Beijing, China
Institute of Zoology, Ukrainian
National Academy of Sciences,
Kiev, Ukraine
Natural History Museum of Los
Angeles County, Los Angeles,
California, U.S.A.
Museo Civico di Storia Naturale
“Giacomo Doria”, Genoa, Italy
Museum of Comparative Zool-
ogy, Harvard University, Cam-
bridge, Mass., U.S.A.
Muséum d Histoire Naturelle,
Geneva, Switzerland
ANIC
BMHH
BMNH
CASC
IZAS
IZUA
LACM
MCSN
MCZC
MHNG
385
Museum of Nature and Human
Activities, Sanda, Hyogo, Japan
Muséum National d’ Histoire
Naturelle, Paris, France
Museum fiir Naturkunde der
Humboldt- Universitit zu Berlin,
Berlin, Germany
National Arthropod Collection,
Mount Albert Research Center,
Auckland, New Zealand
Naturhistorisches Museum, Ba-
sel, Switzerland
Naturhistorisches Museum, Vi-
enna, Austria
Naturhistoriska
Stockholm, Sweden
Oxford University Museum, Ox-
ford, U.K.
South African Museum, Cape
Town, South Africa
National Museum of Natural
History, Washington, D.C.,
U.S.A.
MEASUREMENTS AND INDICES
Head Width (HW). Maximum width of
head, in full-face view, excluding the eyes
(Fig. 1).
Head Width Including 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 mandibles
(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 margin
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 margin (if the anterior margin lacks
any teeth).
Cephalic Index (CI). HW/HL xX 100.
Scape Length (SL). Length of the an-
tennal 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.
MNHA
MNHN
MNHU
NACA
NHMB
NHMV
NHPS Piksmuseet,
OXUM
SAMC
USNM
386
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
S)
Figures 1-3. Measurements illustrated for this study (worker of Pristomyrmex longispinus sp. n.). 1: Head, full-face view; 2:
Entire body, lateral view; 3: Dorsal view of alitrunk, petiole, and postpetiole.
Pronotal Width (PW). Maximum width
of the pronotum in dorsal view (Fig. 3).
Alitrunk 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). Til + 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.) TL1: 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 (PSL1). 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).
Postpetiole Index (PPI). PPW/PPL xX }
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 Taylor's
(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 Pristomyrmex, when erected |
by Mayr (1866), contained one species, P. |
pungens Mayr (=a junior synonym oe Je
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
punctatus (F. Smith)), a member of the
present punctatus group. At that time,
Mayr did not realize that five species de-
scribed by F. Smith (1858, 1860, 1861,
1863, 1865), that is, Myrmica fuscipennis,
Myrmica punctata, Myrmica_ trachylissa,
Myrmica umbripennis, and Solenopsis lae-
vis, also belonged in the new genus. Mayr
(1866) provided a description of the genus,
which actually was simply 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 Pristomyrmex
expanded for the fist time when Mayr
(1886) transferred Myrmica trachylissa F.
Smith to Pristomyrmex. Pristomyrmex tra-
chylissus is now a member of the quadri-
dens group. After that, many species of the
quadridens group were discovered (Bol-
ton, 1981; Donisthorpe, 1949c; Emery,
1887, 1895, 1897, 1900; Forel, 1914; Ka-
ravaiev, 1931, 1933; Taylor, 1965, 1968),
and several more names were added to the
group when Odontomyrmex André and
Hylidris 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-
myrmex.
The members of the third species group
(i.e., umbripennis group) of Pristomyrmex
were recognized first by Emery. He de-
scribed a new species (Pristomyrmex pic-
teti) in 1893 and transferred a species
(Myrmica fuscipennis F. Smith) to Pristo-
myrmex in 1901. Donisthorpe expanded
the umbripennis group: He _ transferred
Myrmica umbripennis F. Smith and Sole-
nopsis laevis F. Smith to Pristomyrmex and
described Pristomyrmex pollux and Pris-
tomyrmex reticulatus (1932, 1946, 1949a).
When Emery (1897) described Pristo-
myrmex coggii, Pristomyrmex levigatus,
and Bristom yrmex lucidus, representatives
of the present levigatus group were added
387
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. Type species: Pristo-
myrmex pungens Mayr, op. cit.: 904 [=Myrmica
punctata F. Smith, 1860: 108; = Pristomyrmex
punctatus (F. Smith) ]; by monotypy.
Odontomyrmex André, 1905: 207. Type species:
Odontomyrmex quadridentatus André, op. cit.:
208; by monotypy. [As a subgenus, thus synonym,
of Pristomyrmex by Forel, 1915: 53.]
Hylidris Weber, 1941: 190. Type species: Hylidris
myersi Weber, loc. cit. (=Pristomyrmex africanus
Karavaiev); by original designation. [Synonymy by
Brown, 1953: 9.]
Dodous Donisthorpe, 1946: 145. Type species: Do-
dous trispinosus 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 from 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) monomorphic myrmicine ants.
*2. Mandible somewhat subtriangular;
masticatory margin of mandible with he ee
to five teeth, Shieh 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
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. bispinosus and P. trispino-
Sus, OF
the apical + the preapical + a lon-
ger diastema + a small denticle + a
shorter diastema (sometimes the
second diastema is indistinct) + a
small basal denticle, as in PR. browni,
or
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
quardridens group, or
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.
trachylissus.
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. divisus 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 three pairs of
lateral denticles (or crenulate shapes) but
sometimes the median tooth rudimentary
(as in some species of the levigatus group)
and sometimes anterior clypeal margin
lacking any distinct denticles (as in P. pro-
fundus, P. divisus, and P. pulcher).
6. Ventral surface of clypeus with a me-
(5)
(6)
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
dian tooth or two lateral teeth, or with a
transverse ridge, or without any ridge or
tooth.
*7. Dorsal labrum with a raised trans-
verse ridge or a few toothlike prominenc-
es, present on the anterior portion of la-
brum in most species.
8. Palp formula l,2, 1372/20 253)4)3 0%
53.
9. Frontal lobes absent in punctatus
and trispinosus groups or weak, as in lev-
igatus, profundus, and quadridens groups,
or somewhat expanded, as in umbripennis
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 umbripennis 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 approximately 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. Alitrunk 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-
nosus and P. browni.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
18. Propodeum armed with a pair of
teeth or spines, except in P. inermis.
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 sometimes without a
ventral process, sometimes 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 rugoreticulum, or
regular striate sculpture. Gaster unsculp-
tured.
28. Dorsal surfaces of head and alitrunk
usually with numerous 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; sometimes
petiole and postpetiole lacking hairs. First
gastral tergite usually without hairs or with
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
389
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. mandibularis), 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 pronotum is armed 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, Mf2+3 usually reduced to vestigial
lines distally; cross-vein m-cu 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); 1r 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-
tomyrmex, is rather stable, with only slight
variations within the different species. For
example, on the forewings, Mf2+3, some-
times becomes an almost entirely vestigial
line, but sometimes it is distinct and rather
long; Rsf4 + Rsf5 is rather thick and long
in some larger species but thin and short
in some smaller species).
390
5
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
7
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. quadridentatus, P. sulcatus, P. browni, P. trispinosus, P. obesus, P.
levigatus, P. picteti, P. pollux, P. umbripennis, and P. punctatus.]
Ergatoid Queen. General characters, in-
cluding the pronotal prominences 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. africanus); 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 smaller 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 margin of labia broadly
concave at center; Mors 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
maximum length of eye; of the other 11
funicular segments, the first segment
shortest, the apical segment longest, the
remaining nine segments much longer
than their broad.
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 completely
exposed in full-face view.
12. Palp formula as in the conspecific
worker in seven species examined (i.e., P.
punctatus, P. quadridens, P. curvulus, P.
brevispinosus, P. sulcatus, 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
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 armaments.
17. Mesonotum well developed, con-
sisting of a large mesoscutum, a rather
large mesoscutellum, and two small axillae.
Notauli usually distinct, forming a Y shape,
but sometimes 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,
overhung by mesoscutellum.
19. Propodeum showing a sloping dor-
sal surface; propodeal armaments 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 sometimes bearing a
small tooth.
25. Positions of spiracles on propo-
deum, petiole, postpetiole, and first gastral
segment similar to those in the conspecific
worker and queen.
narrow,
391
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 small, toothlike, not
meeting
Petiole: With a long anterior peduncle
Forewing: Without m-cu cross-vein;
marginal cell open
Acanthomyrmex
Antennae: 13 segments
Mandibles: Subtriangular, with six to
eight teeth, meeting when they are
closed
Petiole: Similar to that of Pristomyrmex
Forewing: PWith 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
Wheeler’s (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 curved 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 NAMES
WITH SYNONYMIES
(Currently valid names are in boldface)
acerosus: sp. n.
africanus: Pristomyrmex africanus
Karavaiev
=beni
=mbomu
=myersi
=primus
aruensis: Pristomyrmex quadridens var.
aruensis Karavaiev
=quadridens
beni: Hylidris myersi subsp. beni Weber
=africanus
bicolor: stat. n.: Pristomyrmex trachy-
lissa var. bicolor Emery
=taurus syn. n.
bispinosus: Dodous bispinosus Donis-
thorpe
boltoni: sp. n.
brevispinosus: Pristomyrmex brevi-
spinosus Emery
=yaeyamensis syn. n.
browni: sp. n.
castaneicolor: Pristomyrmex castaneico-
lor Donisthorpe
=umbripennis
castor: Pristomyrmex castor Donis-
thorpe
=umbripennis
coggit: Pristomyrmex coggti 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
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.
obesus
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 393
=pegasus syn. n.
occultus: sp. n.
orbiceps: Xiphomyrmex orbiceps Sants-
chi
=laevigatus
orbiculatus: Pristomyrmex orbiculatus
Donisthorpe
=quadridens
parumpunctatus: Pristomyrmex parum-
punctatus Emery
=umbripennis
parvispina: Pristomyrmex parvispina
Emery, 1900: 678. Nomen nu-
dum
pegasus: Pristomyrmex pegasus Mann
=obesus
picteti: Pristomyrmex picteti Emery
=tingiana syn. n.
pollux: Pristomyrmex pollux Donis-
thorpe
primus: Hylidris myersi subsp. primus
Weber
=africanus
profundus: sp. n.
pulcher: sp. n.
punctatus: Myrmica 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 André
=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: Pristomyrmex picteti var. tin-
giana Stitz
=picteti
trachylissus: Myrmica trachylissa_ F.
Smith
trispinosus: Dodous trispinosus Donis-
thorpe
trogor: Pristomyrmex trogor Bolton
umbripennis: Myrmica umbripennis F.
Smith
=castaneicolor syn. n.
=castor syn. n.
=laevis syn. n.
=parumpunctatus 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.
Tle Dorsum of alitrunk in profile not arched,
with mesonotum much higher than
propodeal dorsum, that is, a vertical
cliff present between mesonotum
and propodeal dorsum (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 lamella (Fig. 80)
(profundus group; Asia: Sabah) —-----
Siena ees profundus (p. 515)
Dorsum of alitrunk in profile, excluding
armaments, 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.
SHS 2)) ie aE ae ae ce es 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 ORI PAL AUS AFR IOI
punctatus group P. divisus sp. n. oF
P. fossulatus (Forel) +
P. pulcher sp. n. +
*P. punctatus (F. Smith) + +
rigidus sp. n. *
cribrarius group cribrarius Arnold +
quadridens group africanus Karavaiev +
bicolor Emery +f
brevispinosus Emery ate
collinus sp. n. +
costatus sp. 0. aie
curvulus sp. n. is
eduardi Forel +f
erythropygus Taylor 4p
flatus sp. n. +f
foveolatus Taylor 2s
hirsutus sp. n. a
longispinus sp. n. ati
modestus sp. n. a5
+
+
nitidissimus Donisthorpe
occultus sp. n.
orbiceps (Santschi) oo
quadridens Emery +
quadridentatus (André) +
quindentatus sp. n.
sulcatus Emery
thoracicus Taylor +
trachylissus (F. Smith) +
trogor Bolton +
wheeleri Taylor a
wilsoni Taylor 4s
bispinosus (Donisthorpe)
browni sp. n.
trispinosus (Donisthorpe)
acerosus sp. n.
boltoni sp. n.
coggit Emery
inermis sp. Nn.
largus sp. n.
levigatus Emery
longus sp. n.
lucidus Emery
> mandibularis Mann
*P. minusculus sp. n.
P. obesus Mann
P. simplex sp. n.
profundus group P. profundus sp. n.
umbripennis group P. fuscipennis (F. Smith)
P. picteti Emery
P. pollux Donisthorpe
P. reticulatus Donisthorpe
P. umbripennis (F. Smith)
neh as) ash awlas) mshi as) as) eh melas) ae) ae) is) as)! Se) fas) das) As) taehlac) tas) as) las) be)
N
AS ae) AS} aS) gs) as) Bs) as) else) As) As) ashe
trispinosus group
+++
levigatus group
S+ttt+t+++++¢¢¢¢¢4¢4¢4¢4+
+
Total number of species 1 vl 5 3
Total number of the
endemic species 36 6 5 3
Notes: ORI, PAL, AUS, AFR, and IOI 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 that P. punctatus and P. minusculus occur in
the two regions, respectively.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 395
25
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. pollux 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 (André); 20: P. wheeleri Taylor; 21: P. longispinus sp. n.; 22: P. trispinosus (Donisthorpe); 23: P. browni sp.
n.; 24: P. bicolor Emery; 25: P. wilsoni Taylor. Figure 26. Propodeal spines of the worker of Pristomyrmex browni sp. n., dorsal
view.
396 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
—~
— eee es
——_—>> See
eee Ae aes
34
35 36
—— a
| ——_
37 : ; 2 :
SP
38 39
Figures 27-39. Mandibles of Pristomyrmex workers. 27: P. profundus sp. n.; 28: P. levigatus Emery; 29: P. mandibularis Mann;
30: P. picteti Emery; 31: P. quadridens Emery; 32: P. quindentatus sp. n.; 33: P. quadridentatus (André); 34: P. trachylissus (F.
Smith); 35: P. rigidus sp. n.; 36: P. punctatus (F. Smith); 37: P. browni sp. n.; 38: P. trispinosus (Donisthorpe); 39: P. bispinosus
(Donisthorpe).
QD. Masticatory margin of mandible with New Guinea; Indonesia; Pohnpei
four teeth; the third tooth, counting Is.; Palau Is.; Yap I.; Tonga Is.; Wal-
from the apex, smallest, distinctly lis Is.; found rarely in N. Queens-
smaller than the basal one; diastema land, Australia) _. minusculus (p. 507)
absent between the preapical and Pronotummunanned (Hig). 5
the third tooth (Figs. 28-29) (levi- 5, Postpetiole in profile with an arched an-
gatus group; Asia, Australia) 3 terior face and a steeply sloping pos-
Masticatory margin of mandible with terior face and the apex of postpe-
three to five teeth; if four teeth pre-
sent, then the third tooth, counting
from the apex, similar in size to the
basal one; diastema either present
or indistinct between the preapical
and the third tooth (Figs. 30-39) _ 14
oy Propodeum unarmed (Fig. 10). Petiole
node in profile wedge-shaped (Fig.
45) (Asia: Papua New Guinea) --------
a ee re Lr et oe 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
4. Pronotum armed with a pair of teeth
(Fig. 11) (Asia and Pacific Is.: Papua
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. 40-42, 44). Head narrower;
Dea IN Varco lel 010 Veena ae nk SO ee
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 397
Figures 40-58. Petiole nodes and postpetioles of Pristomyrmex workers. 40A: Dorsal surface of the petiole node of P. obesus
Mann, dorsal view; 41A: Dorsal surface of the petiole node of P. /Jongus sp. n., dorsal view; 40B, 41B, 42-58: Petiole nodes
and postpetioles, lateral view: 40B: P. obesus Mann; 41B: 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. flatus 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
8. Eyes smaller, with two to three omma-
tidia in the longest row (Asia: New
(@uin'ea)) eee boltoni (p. 492)
Eyes larger, usually with five to seven
ommatidia in the longest row ___. 9
9. Larger species, with HW 0.90-0.96 and
cy ZEN HL 0.90-0.90 (Pacific Is.: Pohnpei
59 Zany Pa LT he iia eRe eee largus (p. 497)
Smaller species; HW < 0.80, HL < 0.80
Figures 59-60. Gasters of Pristomyrmex workers, lateral 10 Cea ene Sone hol ae )e “th al Sr rt
view. 59: Gaster of P. simplex sp. n.; 60: Gaster of P. obesus ; PEVOle witht ay Ong
Mann. pinlike process (Fig. 42) (Pacific Is.:
New Hebrides) acerosus (p. 491)
Ventral surface of petiole without a long
6. Dorsum of petiole node laterally com- pinlike process (Figs. 40, 41, 44) — 11
pressed and in dorsal view distinctly Le Each side of petiole with a longitudinal
longer than broad (Fig. 41A) (Asia: carina that separates the tergite
* New Fat oe ranges ‘ 502) from the sternite (Fig. 40). Basal
orsum of petiole node not laterally Ries maya eee
compe eee and in dorsal view Lara cecny on He ube
about as broad as or broader than
long, not longer than broad (Fig.
A TAN) fap sateen aie eas 220 tS, aE
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-
cific Is.: Papua New Guinea, Solo-
mon Is., Nama Is., New Britain Is.)
5 LOE fo, Se) Neel 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)
(Raxerinv® 105.9 VEMD) een tet nate s
“
~“
imperromtale camin ae) ees 8 : :
Dorsum’ of alitrumk with,some scattered’ et ee mandibularis (p. 505)
foveolate punctures. Dorsum of 12(7). Entire first gastral tergite evenly clothed
head between frontal carinae with with numerous erect or suberect
scattered foveolate punctures or fo- hairs (Fig. 60) (Pacific Is.: Solomon
veolate-reticulate sculpture (Figs. LSS) Pea ee ooh eee obesus (p. 509)
(OGY i a ae ge en SAN ee 12 Only a few hairs present usually near the
o>
CS
ay
pas:
SS
L)
S V2
\e
I\N
)
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 antennal fossae and on the genae; 67: Regular striations on the entire dorsum of the head.
13.
14(5
15.
16.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
base of the first gastral tergite (Fig.
BIO) Pe eS ROE PS ae ae Eee
Dorsum of head, except for the scrobes,
with foveolate-reticulate sculpture;
punctures often aligned 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)
13
a ea ees owe Ne 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) ------------
suerte aceite ikabie ioe Ane Ae simplex (p. 512)
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; ein) Poa Poranl a
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, masticatory 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 __.
Eyes larger, usually consisting of 20 or
more ommatidia, containing six to
seven ommatidia in the longest row.
Propodeum with a pair of Roothilike
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-
15
7,
S12) pea aa emus umbripennis (p. 525)
Eyes smaller, 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
[SING ceo sine eda te en oo ee
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)
Propodeal spines shorter and not strong-
16
pollux (p. 521)
18.
19.
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) -—-
399
wb: 2ca Jui dane Ne ee te 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)
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 ARE and head between frontal
carinae unsculptured and smooth __
Dorsum of alitrunk with well developed
coarse reticulum. |Propodeum
armed with a pair of long spines.
Antennal scapes longer, usuall
>0.78; one-sixth to one-fifth of the
length of the scapes projecting be-
yond the occipital margin. Palp for-
TOGNDU Kee Dy Gis aubeee see ae ak a ned nl oe
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)
18
bo
bo
19
Sa arse to Oe See fossulatus (p. 406)
Lateral portions of clypeus, 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
fey eee re OTe AD RE Ns Nr se once
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
TSS euhheit totes sae ees Ree ee RP Rien
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) —-------
20
21
RRs LIN te Pals ea ACE. 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
71
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 with two minute toothlike promi-
view lacking a deep longitudinal fur- nences. Dorsal alitrunk convex, not
row at the midline (Asia: Malaya) - depressed. Sculpture of the sides of
Pe SOPRE PRE ites. ON uno 2 pulcher (p. 408) pedicel segments more coarse (Asia:
21(19). Two or more pairs of erect hairs present Thailand, Malaya, Sarawak, Sabah,
on the dorsum of petiole node (Fig. Brunei, Sumatra) rigidus (p. 415)
46). Basal margin of mandible al- 22(17). Promesonotal suture present. Propodeal
most straight, without a distinct spines developed and long, in dorsal
tooth (Fig. 31). Ventral surface of view joining together at base and
clypeus lacking toothlike promi- forming a “fork” (Fig. 26). Alitrunk
nences. Dorsal alitrunk more or less in profile with a convex promeson-
depressed, with marginate sides. otum and a deeply concave propo-
Sculpture of the sides of pedicel deal dorsum (Figs. 22, 23). Dorsum
segments lighter and finer (wide- of head, at least on the genae and
spread in the east and south of Asia; around the antennal sockets, with
occasionally intercepted at entry regular striate sculpture, lacking fo-
ports in North America) ———----- veolate punctures or rugoreticulum
eae punctatus (p. 410) (Figs. 66, 67) (trispinosus group; In-
A pair of hairs present on the dorsum of dian Ocean Islands) _.... 23
petiole node (Fig. 47). Basal margin Promesonotal suture absent. Propodeal
of mandible with an acute or broad- armaments in dorsal view usually
based triangular tooth (Fig. 35). well separated at the base and not
Ventral surface of clypeus usually resembling a fork. In rare case,
where the propodeal spines are set
close together at the 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 rugoreticulum, but never
showing regular striate sculpture . 25
23. Propodeal spines in dorsal view diver-
gent, in profile almost straight.
Figures 73-75. Lateral portions of the clypei of Pristomyrmex Larger species with HW > 1.00, HL
workers in front of antennal fossae. 73: P. divisus sp. n.; 74: eNO ESI, 2 SO ans Ane 24
P. pulcher sp. n.; 75: P. punctatus (F. Smith). Propodeal spines in dorsal view subpar-
24.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
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 [.)
Dorsum of head and alitrunk entirely
covered with regular long coarse
striations (Fig. 67). Mesonotum with
a pair of strong, blunt digitlike
prominences (Fig. 22) (Indian
Ocean Is.: Mauritius)
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) ---------------------
25(22). Sides of postpetiole with several coarse
26.
longitudinal rugae (Fig. 48). In pro-
file view, the posterodorsal and pos-
teroventral corners of the petiole
node right-angled. Palp formula 4,3
(cribrarius group; Africa: Mozam-
bique, South Africa)
browni (p. 485)
trispinosus (p. 488)
bispinosus (p. 484)
ae cele eat Maree ate 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-
tralia, Africa)
Mandibular dentition arranged as an api-
cal tooth + a preapical + 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 masticatory margin
covered by the three small denticles
is slightly longer than that of diaste-
ma. Pronotum either unarmed or
armed with a pair of short triangular
spines that are shorter than the pro-
jexolenll sormnes eo
Mandibular dentition arranged as an api-
cal + a preapical + a diastema +
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
2)
bo
“I
6
Nw
~“
30.
31.
an apical + a preapical + 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)
Pronotum unarmed. Eyes smaller, with
three ommatidia in the longest row
(Asia: Sumatra)
Pronotum armed with a pair of short tri-
angular spines. Eyes larger, usually
with five to six (rarely with four) om-
matidia in the longest row
Dorsal surfaces of head and alitrunk only
with scattered shallow foveolate
punctures; dorsum of alitrunk with
a smooth and unsculptured median
longitudinal strip (Asia: Indonesia)
Dorsal surfaces of head, except for scro-
bal areas, and alitrunk entirely cov-
ered with well developed coarse ru-
goreticum (Asia: Sarawak, Sabah) --
head between frontal carinae either
smooth or with some scattered fo-
veolate punctures but lacking fove-
olate-reticulate sculpture or rugore-
tL GUL (ha aes eee ee eae etree
Dorsal surfaces of both alitrunk and
head with foveolate-reticulate sculp-
tunejor mugoreticulum a ee
Area of dorsal head between frontal ca-
rinae unsculptured and smooth.
Dorsum of alitrunk without foveo-
late spun ctuie Sue ee ee
Dorsal surfaces of both alitrunk and
head between frontal carinae with
some scattered foveolate punctures
Alitrunk, in dorsal view, with a transverse
ridge at the approximate position of
metanotal groove (Fig. 19). Anterior
clypeal margin with three strong
teeth (Fig. 70)
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)
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)
First gastral tergite lacking any suberect
401
eduardi (p. 440)
28
quindentatus (p. 467)
occultus (p. 455)
. Dorsal surfaces of both alitrunk and
30
31
40
34
erythropygus (p. 441)
402
eee
76 77
aE
pe) Oe ys wes
Se
78
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
oe
Figures 76-79. Characters on the ventral surfaces of the clypei of Pristomyrmex workers. 76: A tooth; 77: Two toothlike
prominences; 78: A short ruga; 79: A long transverse ridge.
hairs. Alitrunk in dorsal view without
rugae at the juncture between the
pronotum and the mesonotum _._.. 33
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 (Australia: New South
Wales, SE Queensland)
BY esi NIVEA OE 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)
pect eine ih el quadridentatus (p. 463)
34(31). Pronotum tuberculate, lacking teeth or
spines (Africa: Ivory Coast, Ghana,
Nigeria, Cameroon, Gabon, Congo
and Angola) orbiceps (p. 456)
Pronotum armed with a pair of teeth or
ES] ON DONS a tS ee 35
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
80 81 82
Figures 80-82. Lamella, circling the base of the antennal
scape of the Pristomyrmex worker, absent (80), entire (81), or
with a broad and deep notch (82).
36.
3M.
38(36).
39.
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
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)
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)
Propodeal spines short or moderately
long, much shorter and slender than
pronotal spines (Fig. 21)
Propodeal spines exceptionally long,
subequal in length to or slightly lon-
ger than pronotal spines (Fig. 25)
(Australia: Queensland) ------------------
eS he abe seh ea A Ee wilsoni (p. 481)
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)
39
REVISION OF THE ANT GENUS PRISTOMYRMEX °
40(30). Petiole and postpetiole lacking erect
Ventral surface of
hairs (Fig. 50).
clypeus with two toothlike promi-
nences (Figs. 77) (Africa: Ghana,
Cameroon, Capen! Angola, Kenya,
Zaire, Sudan)
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.
TASH ry oy Seen en UR noe ate a
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.)
africanus (p. 423)
41
fabs 02a pee 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)
evenly distributed, erect or suberect
hairs. Petiole node with a single
evenly blunt-rounded apex (Fig. 56)
(Asia: Philippines) _
First gastral tergite lacking erect hairs.
Peuole node not showing a single
evenly blunt-rounded apex but with
a higher anterodorsal angle than the
posterodorsall(Hige oi) pee
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)
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)
nitidissimus (p. 453)
42(29). First gastral tergite with numerous,
hirsutus (p. 449)
43
trachylissus (p. 474)
44,
45.
A6.
47(45).
49.
Wang 403
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
Propodeal spines long (0.19-0.30), much
longer than the pronotal armaments
(Bigs 16) Spalpstormulal 213 sae 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
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)
Jn. Wee ew ae eae 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) (aera: Queensland)
ey oe en EN thoracicus (p. 473)
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) Nees Sarawak)
Dimi Fra es ah as modestus (p. 452)
Petiole node in profile higher than long,
with the anterodorsall angle disanee
ly elevated above the posterodorsal;
dorsum and sides of petiole node
without foveolate punctures (Fig.
DID) ence ek Ree eS Rael eee 48
Ventral surface of clypeus lacking a cen-
tral prominent tooth but usually
bearing a transverse ruga (Asia: Ma-
laya, Singapore, Sar Ale Borneo,
Sabah, Philippines) _. costatus (p. 434)
Ventral surface of clypeus with a central
prominent tooth (Fig. 76) __- 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) -—---
Soe ese Re 27 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)
404
(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). z
(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 (P. fossulatus) is confined
to South Africa. Pristomyrmex 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.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
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 masticatory
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 im-
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 from 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
MHNG).
(MCZC, BMNH, ANIC, |
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 margins. 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, surpassing
the occipital margin of head by one-sixth
to one-fifth of their length. Eyes large and
prominent, containing § 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. Pronotum 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 with 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 three 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 margin 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. pulcher. 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
Tetramorium (Xiphomyrmex) fossulatum Forel, 1910:
428. Syntype workers, South Africa: Natal, Will
Broak (Wroughton) (MHNG) lexamined].
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
with § 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—
ONSRE RISA Noss (na—s >)
Mandibles smooth and shining. Denti-
tion of the masticatory margin of mandi-
ble: an apical tooth + a preapical tooth +
a long diastema + 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 * Wang
are fused together. Lateral portions of
clypeus reduced to margins, and antennal
fossae reaching the lateral anterior mar-
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-
Final ruga. Frontal lobes absent; thus, the
anal 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. Propodeum 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, aa a rounded dorsum,
in Moral 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—mesonotum (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.
Male. Unknown.
Comments and Discussion. The position
of P. fossulatus within Pristomyrmex is
somewhat complicated. Pristomyrmex fos-
407
sulatus occurs only in South Africa and
shares certain character states with P. or-
biceps, an African member of the quadri-
dens group. The workers of P. fossulatus
are similar to those of P. orbiceps in (1)
the masticatory margin of mandible with a
diastema, (2) the pronotum with a pair of
broad and blunt tubercles, (3) the size of
eyes, (4) the length of propodeal spines,
and (5) hairs on the head and body (e.g.,
two to three pairs of erect hairs along fie
frontal carinae behind the level of anten-
nal insertions, a pair on the occipital cor-
ners, a pair on the mesonotum). But it is
very hard for me to place P. fossulatus into
the quadridens group because this species
has four segments of the maxillary palpi in
the workers. In the 25 members of the
quadridens group, the maxillary palp of 18
species examined, including P. orbiceps, is
one or two segments.
Pristomyrmex fossulatus, however,
shares important similarities with four
Asian species of the punctatus group @P.
divisus, P. pulcher, P. punctatus, and P. rig-
idus). These critical characters, as shown
in the workers, include (1) a high palp for-
mula, (2) the masticatory margin of man-
dible with a diastema after the preapical
tooth, (3) pronotum without teeth or
spines, (4) anterior face of the petiole node
indistinguishable from the upper surface
of its anterior peduncle, and (5) postpeti-
ole in dorsal view much broader than long.
Thus, P. fossulatus is assigned to the punc-
tatus group. P. fossulatus « can be separated
from the four species as follows: In the
workers of P. fossulatus, the propodeal
spines are moderately long; the antennal
scapes, falling into the range 0.56 to 0.61
(SI 81-88), are only close © the occipital
margin of head, when laid on the dorsal
head; the dorsum of the alitrunk has only
scattered foveolate punctures. In P. divi-
sus, P. pulcher, P. punctatus, and P. rigidus,
the propodeal spines are well developed
and long, much longer than the distance
henwecnt their asoe. the antennal scapes,
with the length 0.70 to 0.94 (usually above
0.78) and SI 91-118 (usually above 98),
408
obviously surpass the occipital margin of
head; the dorsum of the alitrunk is covered
fully with a developed rugoreticulum. It is
possible that P. fossulatus evolved from the
P. divisus-P. pulcher-P. punctatus-P. rigidus
ancestor but later was divergent from the
clade consisting of the four Asian species.
It must be pointed out that if P. fossu-
latus is designated as a member of the
quadridens group, the cribrarius group,
also possessing 4,3 palp formula, would
become insignificant. As a result, P. cri-
brarius (another African species) would
fall into the quadridens group.
Pristomyrmex fossulatus differs from
two African species of the quadridens
group (P. orbiceps and P. africanus) in the
workers as follows:
P. fossulatus
Palp formula 4,3
Dorsum of head between frontal cari-
nae with numerous foveolate punc-
tures
Dorsum of alitrunk with some foveolate
punctures
Petiole node in profile wedge-shaped,
with a triangular apex
P. orbiceps
Palp formula 1,3
Dorsum of head between frontal cari-
nae smooth and shining, except for
few punctures bordering frontal cari-
nae
Dorsum of alitrunk smooth and shining
Petiole node in profile relatively mas-
sive, and broad-rounded dorsally
P. fossulatus
Palp formula 4,3
Ventral surface of clypeus with a weak
transverse ruga
Eyes larger, with 8 to 10 ommatidia in
the longest row
Petiole node in profile wedge-shaped,
with a triangular apex, lacking a dis-
tinct concave between the anterior
face of the node and the dorsal sur-
face of the peduncle
Pronotum with a pair of tubercles
Dorsal head behind the level of anten-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
nal insertions with three to four pairs
of hairs; dorsal alitrunk at most with
a pair of hairs
P. africanus
Palp formula 1,3
Ventral surface of clypeus with two
strong toothlike prominences
Eyes smaller, with four to five ommatid-
ia in the longest row
Petiole node in profile not wedge-
shaped; a distinct concave present be-
tween the anterior face of the node
and the dorsal surface of the pedun-
cle
Pronotum usually with a pair of teeth or
spines but sometimes with a pair of
tubercles
Dorsal head behind the level of anten-
nal insertions with numerous (at least
10 pairs of) hairs; dorsal alitrunk with
several pairs of hairs
Distribution. This species was previous-
ly known only from the type series col-
lected from South Africa (Bolton, 1981).
Two more specimens (ANIC, MCZC) ex-
amined here add the following records:
South Africa: Cape Province, Grahams-
town, Old P. E. Road (L. H. Weatherill).
Ecological Information. Unknown.
Pristomyrmex pulcher sp. n.
Figures 87-88
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 surfaces of —
head and alitrunk, as well as the sides of
pronotum, with well-developed coarse ru-
goreticulum.
Holotype Worker (BMNH). TL 2.96, |
HL 0.76, HW 0.72, CI 95, SL 0.82, ST 114, |
EL 0.20, PW 0.53, AL 0.76. Paratypes, 11 |
workers (BMNH, MCZC, ANIC).
Worker. TL 2.70-3.04, HL 0.70-0.77, |
HW 0.69-0.77, CI 95-100, SL 0.76-0.84, |
ST 103-114, EL 0.19-0.21, PW 0.50-0.54, |
AL 0.70-0.78, PPW 0.25—-0.29, PPL 0.17- |
0.20, PPI 135-156 (n = 11).
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 409
Figures 87-88. Pristomyrmex pulcher sp. n. 87A: Worker head, full-face view; 87B: Showing a curved ruga on the ventral
clypeus; 88: Worker, lateral view.
Mandibles usually with a few longitudi-
nal rugae but smooth near the masticatory
margin. Dentition of the masticatory mar-
gin of mandible: the strongest apical tooth
+ the second strongest preapical + a long
diastema + a broad basal tooth usually
fused by two small denticles. A broad-
based triangular short tooth present about
midway on the basal margin of the man-
dible. Clypeus with sculpture consisting of
a strong median longitudinal carina, a
transverse carina (sometimes curved or
weak or broken), and a few additional
short carinae. Anterior clypeal margin
lacking distinct denticles. Lateral portions
of clypeus, in front of antennal fossae, de-
veloped, not reduced to narrow margins.
Ventral surface of clypeus usually with a
somewhat “M” ruga that is sometimes in-
terrupted at the middle. Palp formula 5,3.
Frontal carinae strong, extending to the
level of the posterior margins of eyes.
Slightly impressed scrobal areas present
lateral to the frontal carinae. Frontal lobes
nearly absent; thus, the antennal articula-
tions are completely exposed. Antennal
scapes, when lying on the dorsal head, sur-
passing the occipital margin of head by
one-sixth to one-fifth of their length. Eyes
large and prominent, containing seven to
nine ommatidia in the longest row. Occip-
ital margin straight or feebly concave. In
dorsal view, dorsum of the alitrunk mar-
ginated and rather depressed. Pronotum
unarmed. Propodeal spines well devel-
oped, acute, and long, much longer than
the distance between their bases. Meta-
pleural lobes triangular. Pedicel segments
as in Figure 88. Dorsum of petiole pedun-
cle, together with the anterior face of the
node, forming a long declivity that reaches
the apex of the node. Ventral surface of
petiole without any appendages or with
only a very narrow rim. Postpetiole in pro-
file convex dorsally, in dorsal view trans-
verse-rectangular and much broader than
long. Dorsal surfaces of head and alitrunk,
as well as the sides of pronotum, with de-
veloped coarse rugoreticulum. Sides of the
rest of the alitrunk irregularly rugulose.
Antennal scapes with a few longitudinal
rugae. Sides of both petiole and postpeti-
ole usually with a few coarse longitudinal
rugae. In dorsal view, petiole and postpe-
tiole each bounded by a rim; dorsal sur-
faces of petiole and postpetiole, except for
rims, very smooth and polished. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with numerous erect or suberect
hairs. A pair of similar hairs present bilat-
erally near the top of petiole and two pairs
usually on the dorsal postpetiole as in Fig-
ure 88. Two or three pairs of long, for-
ward-projecting hairs present near the an-
terior clypeal margin. Scapes and tibiae
with some scattered erect or suberect
410
hairs. Gaster lacking erect or suberect
hairs. Femurs, tibiae light-yellow, in con-
trast with head antennae, alitrunk, tarsi,
pedicel segments, and gaster, which are
reddish-brown.
Queen and Male. Unknown.
Comments. Although P. pulcher, an Ori-
ental species, is very similar, in appearance
of the workers, to P. punctatus, the two
species are in fact not closely related with-
in the punctatus group. Pristomyrmex
pulcher, together with P. divisus, consti-
tutes a clade that is the sister group of the
clade formed by the species P. rigidus and
P. punctatus. The separation of the two
clades is summarized under P. divisus. Ad-
ditional characters separating the workers
of P. pulcher from those of P. punctatus
are as follows:
P. pulcher
A broad, short tooth present on the bas-
al margin of mandible
Dorsum a petiole node with only a pair
of erect hairs
Leg bicolored, with femur and _ tibia
light-yellow and tarsus reddish-brown
Sculpture on the dorsal surfaces of head
and alitrunk, and the sides of petiole
and postpetiole relatively coarse
Propodeal spines shorter, more robust,
and slightly bent basally
P. punctatus
Tooth of the basal margin absent or in-
conspicuous
Dorsum of petiole node with two or
more pairs of hairs
Leg uniformly colored, light-red or red-
teie brown
Sculpture on the dorsal surfaces of head
and alitrunk, and the sides of petiole
and postpetiole relatively fine
Propodeal spines relatively slender, and
straight
The separation of P. pulcher from P. div-
isus is discussed under P. divisus.
Holotype Worker. Malaysia: Negri Sem-
bilan, Pasoh For. Res., primary forest, lit-
ter sample, 3.iv.1994 (L. Ficken).
Paratypes. Five workers with same data
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
as holotype; three workers, Malaysia: Ne-
gri Sembilan, Pasoh For. Res., litter sam-
ple, 3.iv.1994 (M. Brendell, K. Jackson and
L. Ficken); three workers, Malaysia: Neg.
Sembilan, Pasoh For. Res., litter sample,
xi.1994 (M. Brendell, K. Jackson, and S.
Lewis).
Ecological Information. This species has
been collected from a little sample taken
in primary forest.
Pristomyrmex punctatus (F. Smith)
Figures 89-93
Myrmica punctata F. Smith,
workers, Indonesia: Bachian I.
(OXUM) [examined].
Pristomyrmex punctatus (F. Smith) Mayr, 1886: 361.
Pristomyrmex pungens Mayr, 1866: 904. Holotype
worker, Malaysia: Malacca (?) (NHPS) [examined].
Syn. n.
Pristomyrmex japonicus Forel, 1900: 268. Syntype
workers, Japan: Osaka (K. Yamada) (MHNG) [ex-
amined]. [Synonymy by Viehmeyer, 1922: 207].
1860: 108. Syntype
(A. R. Wallace)
Diagnosis (Worker). Eyes with eight or
more ommatidia in the longest row; pron-
otum unarmed; propodeal spines straight,
and long, much longer than the distance
between _ their bees: dorsal surfaces of
both head and alitrunk covered fully with
rugoreticulum; dorsum of petiole node
with two or more pairs of hairs.
Worker wily 2462-32225 Miley 0270201845
HIW 0168-0584 C942 1055S L1057/8=0:S61
SI 102-118, EL 0.15—-0.18, PW 0.48-0.56,
AL 0.70—-0.84, PPW 0.24—0.27, PPL 0.17—
OVA, JAP MAO) Gi —= 70),
Mandibles usually with a few fine lon-
gitudinal rugae but smooth near the mas-
ticatory margin. Dentition of the mastica-
tory margin of mandible: the strongest api-
cal tooth + the second strongest preapical
+ a long diastema + a broad basal tooth
|
'
(or two small basal denticles). Basal mar- —
gin of mandible rather straight, lacking a
Gustine tooth. Clypeus eine leliiee’ more or
less depressed, with a median longitudinal —
carina extending posteriorly through the |
|
frontal area. In some specimens, a few |
short weak rugae present on each side of |
the median carina of the clypeus. Anterior —
clypeal margin equipped with a row of
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
sale
[Z
41]
Figures 89-92. Pristomyrmex punctatus (F. Smith). 89A: Worker head, full-face view; 89B: Showing a curved ruga on the
ventral clypeus; 90: Worker, lateral view; 91: Ergatoid queen head, full-face view; 92: Ergatoid queen, lateral view.
denticles, but sometimes median denticle
indistinct or absent or becoming a broad-
truncated lobe. Lateral portions of clypeus
reduced to margins, so that the antennal
fossae reach the anterior clypeal margin.
Ventral surface of clypeus usually with a
curved ruga as in Figure 89B. Palp for-
mula 5,3. Frontal carinae distinct, extend-
ing to the level of the posterior margins of
eyes. Antennal scrobes weak. Frontal lobes
almost completely absent; thus, the anten-
nal articulations are entirely exposed. An-
tennal scapes, when lying on the dorsal
head, surpassing the occipital margin of
head by one-sixth to one-fifth of their
length. Eyes large, with eight or more om-
matidia in the longest row. Occipital mar-
gin feebly concave. Dorsum of the alitrunk
in dorsal view marginated and more or less
depressed (Fig. 93). Pronotum unarmed.
Propodeal spines long, acute, but slender.
Metapleural lobes dentiform and acute.
Petiole node in profile wedge-shaped, with
a triangular apex. Postpetiole in profile
convex dorsally; in dorsal view transverse-
rectangular, much broader than long, and
also broader than the petiole. Dorsal sur-
faces of head and alitrunk as well as sides
of pronotum covered fully with rugoreti-
culum, but scrobal areas with only several
transverse rugae. Sides of the rest of the
alitrunk with numerous irregular rugae.
Sides of petiole and postpetiole usually
with a few fine longitudinal rugae; some-
times rugae very weak and broken. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with numerous erect to suberect
long hairs. Two (or more) pairs of hairs
412
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figure 93. Showing that the worker of Pristomyrmex punctatus (F. Smith) has a more or less depressed dorsum of alitrunk.
present, respectively, bilaterally on the
dorsums of petiole node and postpetiole,
of which usually a pair shorter and the oth-
er pair longer. First gastral tergite lacking
erect or suberect hairs. A few pairs of long,
forward-projecting hairs present near the
anterior clypeal margin that are symmet-
rical on the two sides of the midpoint.
Scapes and tibiae with numerous erect to
suberect short hairs. Color reddish-brown,
but sometimes the gaster much darker or
the appendages slightly lighter.
Ergatoid Queen. ae, 3.60. 3972: FUE
0.86, 0.88: HW 0.94, 0.94; CI ude 109: SL
0.89, 0.91; SI 97, 97; EL 0.23, 0.23: PW
0.66, 0.68; AL 0.94, 0.96; PPW. 0.31, 0.32
PPE 0.23) 0324- PRT T3335) G7 = 2):
Closely resembling the worker in the
structure of mandibles, clypeus, petiole,
postpetiole and gaster and also in sculp-
ture, color, and pilosity. But the head with
three ocelli; eyes larger; pronotum and
propodeum narrower than those of work-
er; Mesonotum more convex; an impres-
sion present at the approximate positions
of promesonotal suture and of metanotal
groove, respectively; propodeal spines
stronger than in worker. Wing absent, but
the rudimentary vestige of a wing is pres-
ent on the each lateral margin of the me-
sonotum.
Queen. Unions
Male. TL 3.22, HL 0.60, HW 0.57, CI
95) SEAOSs Sie 325 EIWilis OO eerie, 0.32.
PW 0.74, AL 1.04, PPW 0.26, PPL 0.17,
PPI 153 (n = 1; one specimen [MCZC]
collected from Nara, Japan, by Silvestri on
July 21, 1925, was examined).
Head, including the eyes, broader than
long. Clypeus transverse, with a median
short carina. Frontal area with a median
longitudinal carina. Frontal carinae short,
slightly beyond the posterior margins of —
antennal sockets. Palp formula 5,3. On the
mesoscutum, notauli pronounced, forming
a Y shape; parapsidal furrows superficially _
impressed. Propodeum with a pair of
teeth. Metapleural lobes subtriangular.
Middle and hind tibiae each with a simple
spur. Petiole node wedge-shaped, with a
triangular apex; dorsum ae petiole pedun-
cle a ming a declivity that reaches the top
of the monies Postpetiole i in profile rounded
dorsally, in dorsal view transverse-rectan-
gular and distinctly broader than long.
Dorsum of head smooth and shining, ex-
cept for few short rugae present behind —
the posterior margin of clypeus. Pronotum —
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
and mesoscutum smooth, except for those
marked sutures, but mesoscutellum and
propodeum sculptured with several longi-
tudinal rugae. Sides of petiole with a few
rugae. All dorsal surfaces with abundant
erect or suberect hairs, but hairs on the
legs and on the scapes somewhat ap-
pressed. Colour reddish-brown; wing
light-yellow.
Comments and Discussion. The separa-
tion of P. punctatus from both P. divisus
and P. pulcher is summarized in the dis-
cussions of both P. divisus and P. pulcher.
The following characters can be used to
separate the workers of P. punctatus from
those of its closest relative, P. rigidus:
P. punctatus
Dorsum of petiole node with two or
more pairs of hairs
Tooth absent or inconspicuous from
basal margin of mandible
Dorsal surfaces of head and _alitrunk,
and the sides of petiole and postpe-
tiole more finely sculptured
Propodeal spines relatively slender
Clypeus with a median longitudinal ca-
rina that meets the anterior clypeal
margin
Dorsum of alitrunk in dorsal view, mar-
ginated, and more or less depressed
Ventral surface of clypeus with a curved
ruga but lacking distinct toothlike
prominences
P. rigidus
Dorsum of petiole node only with a sin-
gle pair of hairs
In type specimens, a strongly prominent
tooth present on basal margin of
mandible
Dorsal surfaces of head and alitrunk,
and the sides of petiole and postpe-
tiole more coarsely sculptured
Propodeal spines relatively robust
Median clypeal carina often not reach-
ing the anterior clypeal margin
Dorsum of alitrunk in dorsal view, con-
vex
Ventral surface of clypeus usually with
two minute toothlike prominences
413
In addition, an ergatoid queen caste is
present in P. punctatus but not seen in P.
rigidus, while a normal queen caste exists
in P. rigidus but has not been found in P.
punctatus.
Without a doubt, P. punctatus originat-
ed in the subtropics or tropics of Asia, as
its three close relatives, P. rigidus, P.
pulcher, and P. divisus, are restricted to
Sumatra, Malaysia, Brunei, Thailand, and
the Philippines, respectively. But P. punc-
tatus has a very large range, from New
Guinea, Indonesia to Malaysia, Thailand,
and then north to China and Japan, indi-
cating its exceptional dispersal ability and
tolerance of temperate climates. Pristo-
myrmex punctatus has shown some ten-
dency for dispersal by humens. Intercep-
tion records from ports in North America
suggest that humen commerce may have
played a role in this species’ spread in tem-
perate Asia.
The chromosome numbers of this spe-
cies, reported by Imai (1966) and Itow et
al. (1984), are 2n (diploid) = 24 in the ce-
rebral ganglion cells of the workers and n
(haploid) = 12 in the spermatocytes of the
males. The larva of this species was de-
scribed by Wheeler and Wheeler (1954).
Biology. It appears that P. punctatus oc-
curs in open habitats. My impression
stems from field experience in China as
well as from data associated with speci-
mens. This species lacks a normal queen
caste; mature colonies usually contain sev-
eral thousand workers and a few males,
but ergatoid queens are rarely found. Eggs
are normally laid by workers and can de-
velop into workers or males. Ergatoid
queens, when present, can also lay eggs.
Brood production begins in April and lasts
until the end of September. Young workers
remain inside the nest and lay eggs. Older
workers forage but lose the ability to lay
eggs. Nests are often constructed in leaf
litter from June to August but in the soil
around trees from September to October.
Sometimes arboreal nests are constructed
on dead standing trees or in partially dead
parts of living trees. The nest entrances of
414
the arboreal nests and those under rotten
wood are often covered with soil particles.
See the following references for informa-
tion on these and other aspects of the bi-
ology of P. punctatus (Itow et al., 1984;
Mizutani, 1980, 1982; Tsuji, 1988a, 1988b,
1988c, 1990a, 1990b, 1994, 1995; Tsuji
and Itd, 1986).
Material Examined (ANIC, AMNH,
BMHH, BMNH, IZAS, LACM, MCZC,
NHMV, USNM). New Guinea: Trian Jaya
Ransik (Shah); NETH. Vogelkop, Dano-
waria (J. L. Gressitt); Vogelkop, Manok-
wari, 75 m (D. Elmo Hardy); Vogelkop,
Fak Fak, S. coast of Bomberai, 10 to 100
m (J. L. Gressitt). Borneo: Sarawak, Nanga
Pelagus near Kapit, 180 to 585 m (T. C.
Maa); Sarawak, Merirai Valley (T. C. Maa);
Sarawak, Bau, Lake Area (T. C. Maa);
North Borneo, Tawau, Quoin Hill, Cocoa
Res. Sta. (K. J. Kuncheria); North Borneo,
Forest Camp, 19 km N of Kalabakan (K.
J. Kuncheria); North Borneo (SE), Forest
Camp, 9.8 km SW of Tenom (Y. Hirashi-
ma). Java: S. Coast, Sukawayani, 2 m, jun-
gle remnant (J. L. Gressitt). Philippines:
Samar (McGregor); Mindoro, San Jose (E.
S. Ross); P.R. Mindanao, Talacogon,
8.32°N, 125.39°E, relict rainforest, on
Agusan River (B. B. Lowery). Singapore
(H. Overbeck). Thailand: Chiang Mai
Prov., Chiang Rai, Fang Hort. Res. Sta.(D.
G. Furth); Chiang Mai Prov., 18.70°N
98.82°E, Mae Wang Dist., Ban Huai
Thong, 360 m, agricultural area (orchard),
#96-116 (foragers on bamboo shoot) (R. R.
Snelling and Saowapa Sonthichai). Viet-
nam: Perfume Pagoda (P. Jolivet); Cuc
Phuong Forest (P. Jolivet). China: Hainan
Is. (J. L. Gressitt); Guangxi, Xingping (D.
G. Furth); Guangxi, Guilin, Qixing Park
(D. G. Furth); Hong Kong, N.T., Campus
CGaUPAE Ke Shatines 2223 SeNe eA aS atseecar
20 m, #96-6 (R. R. Snelling); Hong Kong,
in bank of mixed orchard (R. Winney); Tai-
po (G. P. Tung); Shatin (Silvestri); Guang-
dong Prov., 60 km W of Guangzhou, Ding-
Hu Mts. (Boucek); Back Liang (S. F.
Light); Fujian Prov., Jiangle (Minsheng
Wang); Yenping (S. Ling); Zhejiang Prov.,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Mokanshan (N. Gist Gee); Shanghai (Sil-
vestri); Soochow (N. Gist Gee); Nanking
(G. P. Tung); Hubei Prov., Xiangfan (Min-
sheng Wang); Guizhou, Leishan (Min-
sheng Wang); Guizhou, Jiangkou (Min-
sheng Wang); Szechwan Prov., Hsinching
A F (W. L. Brown); Szechuen, near Mt.
Ormel (D. C. Graham); Szechuen, Suifu
(D. C. Graham); Szechuen, Jang Chen Pu
near Mt. Omei (D. C. Graham); Taiwan,
Suisha (R. Takahashi); Taiwan, Kosempo
(H. Sauter); Taiwan, Bukai (L. Gressitt);
Taiwan, Taihoku (J. Sonan, T. Shiraki); Tai-
wan, Hassenzan (?); Taiwan, Nantou, Lan
Wa Chu (D. G. Furth); Taiwan, Nantou,
Sung Kang (D. G. Furth); Taiwan, Nantou,
Hueisun (D. G. Furth); Taiwan, Nantou,
Wushe (D. G. Furth); Taiwan, Taipei, Au-
kung (D. G. Furth). Japan: Okinawa, Ku-
nigami (Yonaha-dake), under chips (F. G.
Werner); Ryukyu, Okinawa (S. M. Fuller-
ton); Amami-Oshima, Loochoo Is. (R. Tak-
ahashi); Kagoshima (Silvestri); Kumamotu
(Silvestri); Kyushu, Amakusa, Tomioka (S.
Murakami); Kyushu Is., Klyamacho, Kak-
inohara (D. G. Furth); Kyushu Is., Mt. Hi-
kosan, Soeda Notoge Pass (D. G. Furth);
Kyushu Is., Mt. Hikosan, Biol. Sta. (D. G.
Furth); Fukuoka Pref.,[zuka-machi, Joro-
gahara (D. G. Furth); Shikoku Is., Toku-
shima, Kawamata (M. Azuma); Hyogo, Mt.
Rokko (M. Azuma); Hyogo, Takarazuka,
Namaze (M. Tanaka); 8 mi N. of Kyoto (P.
Hammond); Idzu (S. Akiyama); Kyoto (Sil-
vestri); Mt. Maya (Silvestri); Michino-o |
(Silvestri); Nara (Silvestri); Kuwana (?);
Honshu, Toyama Pref., Toyama city (D. G. |
Furth and K. Suzuki); Nagano Pref., Mat-_
(D. Gs
Furth); Kamakura (H. Nagase); Kanagawa |
Pref., Kamakura (H. Nagase); Kanagawa |
Pref., Odawara (M. Kubota); Tokyo (L. |
Gressitt); Tokyo Pref., Hachioji, Minami- |
Asakawa (D. G. Furth); Yokohama (?);
Chiba Pref., Ichikawa, Konodai (D. G. |
Furth); Bonin Is., Chichi-jima (H. Ikeda); |
Bonin Is., Chichi-jima, Omura, Camp |
sumoto, Shimauchi-Shimoda
beach (F. M. Snyder); Bonin Is., Chichi-
jima, Yatsuse R., Gen.’s beach (F. M. Sny- |
der); Bonin Is., Chichijima, Miyanchama,
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
415
yar RS
Figures 94-97. Pristomyrmex rigidus sp. n. 94A: Worker head, full-face view; 94B: Showing two small denticles on the ventral
clypeus; 95: Worker, lateral view; 96: Queen head, full-face view; 97: Queen, lateral view.
Jack Wm’s beach (F. M. Snyder); Bonin
Is., Chichijima, Sakai-ura, Bull beach (F.
M. Snyder).
I have also examined 11 workers col-
lected at the two entry ports of the United
States (USNM): nine of them, by Harley
and Albrecht, on November 20, 1928,
from Philadelphia, Pennsylvania, in lily
bulbs imported from Japan; the other two
specimens, by J. F. Byrnes, on September
25, 1967, from Anchorage, Alaska, on Ger-
beria sp. imported from Japan. It appears,
however, that P. punctatus has not yet be-
come established in the United States
(Cover, personal communication).
In addition, this species has also been
reported from North Korea and South Ko-
rea (Collingwood, 1976, 1981; Kim and
Kim, 1982: Kim and Kim, 1983).
Pristomyrmex rigidus sp. n.
Figures 94-97
Diagnosis (Worker). Antennal fossae
reaching the anterior clypeal margin; eyes
with six to eight ommatidia in the longest
row; pronotum unarmed; dorsal surfaces
416
of head and alitrunk covered fully with
well-developed coarse rugoreticulum; dor-
sum of petiole node with a pair of hairs.
Holotype Worker (BMNH). TL. 3.40,
HL 0.98, HW 0.94, CI 96, SL 0.91, SI 97,
EL 0.17, PW 0.64, AL 0.86. Paratypes: 11
workers (MHNG, BMNH, MCZC).
Worker. TL 2.73-3.44, HL 0.75—0.98,
HW 0.74—0.94, CI 93-102, SL 0.70—0.94,
SI 91-103, EL 0.14—0.17, PW 0.50_-0.64,
AL 0.70—0.86, PPW 0.24—0.30, PPL 0.16—
O20VPEIMIA4=67 (r= 32):
Mandibles with a few coarse longitudi-
nal rugae that usually do not reach to the
vicinity of the masticatory margin. Denti-
tion of the masticatory margin of mandi-
ble: the strongest apical tooth + the sec-
ond strongest preapical + a long diastema
+ a truncated basal tooth (or two small
denticles). A strongly developed tooth or a
broad, subtriangular short prominence
present about midway on the basal margin
of mandible. Clypeus shield shaped, more
or less depressed, with a median longitu-
dinal carina that usually does not reach to
the anterior clypeal margin; sometimes a
few additional weak rugae are present on
the clypeus. Anterior clypeal margin usu-
ally with a median denticle and three oth-
ers on each side, but sometimes with a lat-
eral denticle indistinct or two lateral small
denticles fused into a larger one. Lateral
portions of clypeus reduced to margins,
and antennal fossae reaching the anterior
clypeal margin. Ventral surface of clypeus
usually with two minute toothlike promi-
nences (Fig. 94B), but sometimes the
prominences are very weak. Palp formula
5,3. Frontal carinae extending to the level
of the posterior margins of eyes. Antennal
scrobes indistinct. Frontal lobes absent, so
that the antennal articulations are entirely
exposed. Antennal scapes, when lying on
the dorsal head, surpassing the occipital
margin of head by one-sixth to one-fifth of
es length. Eyes containing six to eight
Onninatidia in the longest row. Occipital
margin rather straight or slightly concave.
Dorsal surface of alerank serncan lia con-
vex. Pronotum unarmed. Propodeal spines
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
robust, acute, and long, much longer than
the distance between their bases. Meta-
pleural lobes triangular and acute. Pedicel
segments in profile as in Figure 95. In pro-
file, anterior face of the petiole node, to-
gether with the dorsal surface of petiole
peduncle, forming a long declivity that
reaches the top of the node. Petiole node
in profile with an approximately right-an-
gled apex. Dorsum of postpetiole in profile
sometimes angulate but sometimes round-
ed. In dorsal view, postpetiole transverse-
rectangular, much broader than long and
also broader than the petiole node. Dorsal
surfaces of head and alitrunk as well as the
sides of pronotum fully covered with
coarse rugoreticulum. Sides of the rest of
alitrunk with numerous irregular coarse
rugae. Sides of petiole node and postpe-
tiole usually with a few rims as illustrated
in Figure 95, but sometimes some of the
rims rather weak. Petiole and postpetiole
in dorsal view, except for rims, very
smooth and polished. Gaster unsculp-
tured. Dorsal surfaces of head and alitrunk
with numerous erect or suberect long
hairs. A pair of similar hairs present on the
dorsal petiole, and usually two pairs on the
postpetiole as shown in Figure 95. Two or
three pairs of long, forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with numerous
erect or suberect hairs. First gastral tergite
lacking erect or suberect hairs. Color uni-
formly reddish-brown.
Queen. TL 3.30-4.02, HL O;S5=15 OR
HW 0.86—1.10, CI 100-102, SL 0.76—0.88,
SI 80-91, EL 0.18—0.24, PW 0.66—0.86,
AL 0.84—1.08,
0.22, PPI 140-164 (n = 6).
Generally similar to worker, except for
in addition, the sculp- |
ture of mesoscutum is weaker and sparse. |
caste differences:
Male. Unknown.
Comments. Pristomyrmex rigidus occurs
in the Oriental region. It is closely related
to P. punctatus. These two species consti- —
tute a clade. The separation of the two |
species is summarized under P. punctatus. —
Characters separating P. rigidus from the —
PPW 0.28-0.36, PPL 0.20- |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
P. divisus-P. pulcher clade are given under
P. divisus.
Discussion. The material that I have ex-
amined may contain two species. Speci-
mens from Thailand possess an anterodor-
sally angulate postpetiole in profile, a
strongly prominent tooth on the basal mar-
gin of the mandible, and a truncated basal
tooth on the masticatory margin of the
mandible. Specimens from Indonesia, Ma-
laysia, and Brunei have a rounded dorsal
surface of the postpetiole, a weak tooth on
the basal margin of the mandible, and two
small basal denticles on the masticatory
margin. But a single, variable species is
maintained for the present because avail-
able material is still limited. Further col-
lecting and studying are needed.
Holotype Worker. Thailand: 26. Kaeng
Krachan NP., 19.xi.1985 (L6bl and Burcke
hardt).
Paratypes. 11 workers with same data as
holotype.
Additional Material Examined (ANIC,
BMNH, MCZC, MHNG, NHMV). Thai-
land: Khao Sabap NP. (L6bl and Burck-
hardt). Brunei: Ulu Belalong, W. ridge,
500 ft (R. Levy). Malaysia: Sabah, Tawan
Quoin Hill, 750 ft, rainforest, leaf mold,
berlesate (R. W. Taylor); Sabah, Poring
Hot Springs, 500 to 900 m (Burckhardt
and Lébl); Sabah, Poring Hot Springs,
Langanan Falls, 950 m (L6bl and Burck-
hardt); Sabah, Batu Punggul Resort, pri-
mary forest, sifting (P); Sarawak, 4th Di-
vision, Gn. Mulu NP., limestone forest,
200 m, pitfall trap (H. Vallack); Sarawak,
Gn Matang, 20 km W Kuching, 800 m,
Prbimentane forest (L6bl and Bivekhar dt);
Negri Sembilan, Pasoh For. Res., litter,
primary forest (L. Ficken; M. Brendell, K.
Jackson, and L. Ficken); Kuala, Lumpur
(B. Bolton); Upper Gombak Val., near K.
Lumpur, rainforest, berlesate, ca. 1,500 ft
(R. W. Taylor); Kedah State, Gunong Jeral,
5.48N, 100.62E, 550 m, rainforest, berle-
sate (R. W. Taylor and R. A. Barrett); Pa-
hang, Batu Caves N Kuala, Lumpur (L6bl
and Calame); Pahang Gombak, forest litter
(L6bl and Calame). Indonesia: Sumatra,
417
Liwa, 5.04S, 104.03E, rainforest, litter (M.
S. Harvey).
Ecological Information. This species oc-
curs in rainforest and has been taken in
litter berlesates and pitfall traps.
THE CRIBRARIUS GROUP
Worker. Medium sized, with the follow-
ing combination of characters.
(1) Masticatory margin of mandibles
with a long diastema between the preapi-
cal and the basal tooth.
(2) Palp formula 4,3 (Bolton, 1981).
(3) Frontal carinae present.
(4) Lateral portions of clypeus reduced
to a margin, and the antennal fossae reach-
ing the anterior clypeal margin.
(5) Frontal lobes absent.
(6) Lamella that encircles the base of
antennal scape, entire.
(7) Dorsum of alitrunk convex, not de-
pressed; pro-mesonotal suture absent.
(8) Pronotum armed with a pair of
strong, acute, short spines.
(9) Propodeal spines well developed
and long; in dorsal view not forming a
Stork
(10) Petiole node thick in profile; its an-
terodorsal, posterodorsal, and posteroven-
tral corners showing right angles approxi-
mately.
This group contains one species, P. cri-
brarius, from South Africa and Mozam-
bique.
Pristomyrmex cribrarius probably
evolved from an Oriental ancestor of the
punctatus group because P. cribrarius is
similar, in the workers, to some Oriental
species of the punctatus group in the fol-
lowing characters: (1) a high palp formula;
(2) well-developed, long propodeal spines;
and (3) a few coarse longitudinal rugae
present on the petiole and on the post-
petiole, respectively. However, I erect the
cribrarius group to accommodate this spe-
cies instead of placing it in the punctatus
group because it possesses a distinct pet-
ole node in the workers and queens (i.e.,
in profile, the anterodorsal, posterodorsal,
and posteroventral corners of the node
418
right-angled), which is not seen in the oth-
er species groups. In addition, the workers
of P. cribrarius possess a pair of robust
pronotal spines that are absent in the
punctatus group.
Without a higher palp formula (4,3), P.
cribrarius would fall into the quadridens
group. The quadridens group now con-
tains 25 species, but the palp formulae of
18 species examined are 2,3, 2,2, and 1,3.
If P. cribrarius were placed in the quad-
ridens group, the punctatus group (differ-
ing from the quadridens group primarily
by its high palp formula) would have no
grounds to exist, as two species of the
quadridens group, P. eduardi and P. orbi-
ceps, also lack pronotal armaments.
Pristomyrmex cribrarius is also some-
what similar to the three members of the
trispinosus group, from Mauritius. They
all possess robust, acute, but short prono-
tal spines; long, well-developed propodeal
spines; and a very convex dorsum of the
alitrunk in the workers. The similarity,
however, is superficial. Pristomyrmex cri-
brarius differs from the trispinosus group
in the workers in several important char-
acters: The frontal carinae are absent in
the trispinosus group but extend to the
level of the posterior margins of eyes in P.
cribrarius; the promesonotal suture is
present in the trispinosus group but absent
in P. cribrarius; the propodeal spines in
dorsal view are fused together at the base
in the trispinosus group, as opposed to be
separated at the base in P. cribrarius; palp
formula is lower (1,2) in the trispinosus
group but higher (4,3) in P. cribrarius; fo-
veolate punctures on the dorsal surfaces of
the head and the alitrunk are absent in the
trispinosus group but dense in P. cribrar-
ius; and numerous hairs are present on the
first gastral tergite in the trispinosus group
but not seen in P. cribrarius.
Finally, P. cribrarius possesses (1) a long
diastema on the masticatory margin ae
mandible separating the preapical _ tooth
from the basal tooth, (2) a higher palp for-
mula (4,3), and (3) a pair of Tobuet spines
on the pronotum in the workers, showing
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
it is not closely related to the levigatus,
profundus, and umbripennis groups.
Pristomyrmex cribrarius Arnold
Figures 98-101
Pristomyrmex cribrarius Arnold, 1926: 281. Holotype
queen, Mozambique: Amatongas Forest (G. Ar
nold) (SAMC) [examined].
Diagnosis (Worker). See characters 8
and 10 under the group; additional char-
acters including dorsal surfaces of head
and alitrunk with foveolate-reticulate
sculpture and postpetiole with a few
coarse longitudinal rugae on each side.
Worker. TL 3.10-3.54, HL 0.82-0.96,
HW 0.89-1.02, CI 106—110, SL 0.72—0.80,
SI7S—o2" eI, 0.19-0.22, PW 0.55-—0.64,
AlE0FS0=0;905 BPVWV40:30=0S 65 RR 0:20=
OV IAF ISO Go = 2).
Mandibles with a few longitudinal basal
rugae. Masticatory margin of mandible
with an apical tooth + a preapical tooth +
a long diastema + a broad and truncated
basal tooth. Basal margin of mandible lack-
ing a toothlike prominence or curved lobe.
Clypeus with a strong median longitudinal
carina. Anterior clypeal margin wath a me-
dian truncated lobe and usually three to
four denticles on each side. Ventral surface
of clypeus with two weak toothlike prom-
inences, or unarmed. Palp formula 4,3
(Bolton, 1981). Frontal carinae extending
to the level of the posterior margins of —
eyes. Antennal scrobes slightly concave.
Antennal articulations entirely exposed.
Antennal scapes, when lying on the dorsal :
head, not reaching the occipital margin ofl
head. Eyes with 9 to 10 ommatidia in the |
longest row. Profile shape of alitrunk and |
pedicel segments as in Figure 99. Dorsum
of hla in profile, strongly convex.
Pronotum with a pair of robust short |
spines, ca. 0.06 to 0.08. Propodeal spines |
well developed and long, ca. 0.25 to 0.30,
much longer than the distance between
their bases. Metapleural lobes small and |
triangular. Petiole node high and thick in
profile: its anterodorsal, posterodorsal, and |
posteroventral corners showing right an-
gles approximately. Postpetiole high ings
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
419
Figures 98-101.
profile, curved dorsally; in dorsal view,
transverse-rectangular and much broader
than long. Dorsum of head, except for the
scrobal areas, with foveolate-reticulate
sculpture. Dorsum of alitrunk entirely
sculptured with coarse longitudinal rugae
and blunt foveolate punctures between ru-
gae. Sides of alitrunk irregularly rugulose.
Each side of petiole node and postpetiole
with a few coarse longitudinal rugae. Gas-
ter unsculptured. Dorsal surfaces of head
and alitrunk with numerous erect or sub-
erect hairs. Petiole and postpetiole each
with a few pairs of hairs dorsally. First gas-
tral tergite lacking hairs. Anterior clypeal
margin with a row of forward-projecting
hairs. Scapes and tibiae with erect or sub-
Pristomyrmex cribrarius Arnold. 98A: Worker head, full-face view; 98B: Showing two minute denticles on the
ventral clypeus; 99: Worker, lateral view; 100A: Queen head, full-face view; 100B: Showing two minute denticles on the ventral
clypeus; 101: Queen, lateral view.
erect short hairs. Color reddish-brown, but
gaster darker.
Queen. TL 3.84, HL 0.98, HW 1.04, Cl
1065 SE0:805 SI 7 EE O265 BW 0.325
ING WO sy IRA, O86, IPI OLR) IRE lay ~@
= |),
Generally similar to worker, except for
caste differences; in addition, pronotal ar-
maments absent, eyes larger than in con-
specific worker.
Male. Unknown.
Comments and Discussion. At first sight,
P. cribrarius somewhat resembles two
Australian species of the quadridens
group, P. thoracicus and P. foveolatus:
Their workers all possess a pair of short
pronotal spines, a pair of long propodeal
420
spines, foveolate-reticulate sculpture on
the dorsal head, and a transversely broad
postpetiole in dorsal view. However, P. cri-
brarius is rather different from P. thora-
cicus and P. foveolatus. In the workers of
P. cribrarius, palp formula is 4,3; the ven-
tral surface of clypeus shows two minute
teeth; the dorsum of alitrunk is very con-
vex; the petiole and the postpetiole are
sculptured with a few coarse longitudinal
rugae; the eyes are larger, with 8 to 10 om-
matidia in the longest row; and the petiole
node, in profile, shows three right angles.
But in the workers of P. thoracicus and P.
foveolatus, palp formula is 2,3; the ventral
surface of clypeus only has a single distinct
tooth at the center; the dorsum of alitrunk
is somewhat depressed; the petiole and the
postpetiole are unsculptured; the eyes are
smaller, with four to six ommatidia in the
longest row; and the petiole node in pro-
file lacks an acute posterodorsal angle.
In the African Pristomyrmex fauna, P.
cribrarius is easily recognized by the
coarse longitudinal rugae on the dorsal al-
itrunk anak on the des of the petiole and
the postpetiole; its very convex dorsal ali-
trunk; its well-developed, long propodeal
spines; its distinct petiole node; and the
numerous erect hairs on the alitrunk, pet-
iole, and postpetiole.
Distribution. South Africa, Mozambique
(Bolton, 1981).
Ecological Information. Specimens have
been collected by W. L. and D. E. Brown
in South Africa on sand, in coast vine for-
est; by J. C. Faure, in South Africa (Zu-
luland, St Lucia Lake), “by sifting the de-
tritus and damp decaying leaves fond un-
der bushes (Arnold, 1948)” ; and the ho-
lotype by G. Amold on a tree trunk
(Arnold, 1926).
THE QUADRIDENS GROUP
Worker. Usually medium- to large-sized
ants (ble OM/3=1-46. SEW. O16S8=I"623 Ee
2.90-6.48) with the following combination
of characters.
(1) Masticatory margin of mandibles
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
with three to five teeth, which have one of
the following three arrangements:
a. the strongest apical + the second
strongest preapical + a long diaste-
ma + two small teeth of similar size
(or one basal tooth, which is some-
times formed by the fusion of the
two small teeth) or
b. the strongest apical + the second
strongest preapical + a diastema +
three small teeth of similar size or
c. the strongest apical + the second
strongest preapical + an intercalary
small tooth + a very short diastema
(or this diastema indistinct) + two
small teeth of similar size (i.e., as
shown in P. trachylissus).
(2) Anterior clypeal margin with five or
more denticles in most species, but several
species having only three teeth or promi-
nences.
(3) Lateral portions of clypeus in front
of antennal fossae reduced to a margin,
and the antennal fossae reaching the an-
terior clypeal margin.
ae Palp formula 1 - a 11 species), o
2 (three species), or 2,3 (four specail
a Frontal carinae nonre extending to
the level of the posterior margins of eyes,
with the exception in P. ert ythropt ygus, P.
longispinus, P. trogor, and P. wilsoni. |
(6) Frontal lobes indistinct or very
weak. |
(7) Antennal scrobes shallow or absent. |
(8) Lamella, encircling the base of an- —
tennal scape, entire. |
(9) Dorsum of alitrunk lacking pro-me-_
sonotal suture. |
(10) Pronotum armed with small teeth
to well-developed spines, except in P. ed- |
uardi and P. orbiceps.
(11) Petiole node in profile usually.
high, with the anterodorsal angle elevated
above the posterodorsal, but sometimes |
showing other forms. ;
(12) Dorsum of head without sculpture, |
with scattered foveolate punctures, or with —
foveolate-reticulate sculpture or rugoreti- |
culum.
This group currently contains 25 spe-—
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
cies, accounting for almost half the genus.
Of them, three occur in Africa, six in Aus-
tralia and 16 in the Oriental region. In
fact, like many Pristomyrmex, many spe-
cies in this group have a restricted geo-
graphic range.
The quadridens group species are close-
ly related to the cribrarius and punctatus
groups but differ from them, most impor-
tantly by their reduced palp formulae: 1,3,
or 2,2, or 2,3 (4,3 in the cribrarius group
and 4,3 or 5,3 in the punctatus group).
The quadridens group differs from the
trispinosus group because in the workers
of the quadridens group, a promesonotal
suture (or impression) is absent, the dor-
sum of the propodeum in profile is not
deeply concave, the propodeal spines in
dorsal view do not form a fork, and regular
striation is absent from the dorsal surfaces
of the head and the alitrunk.
The quadridens group differs from the
levigatus, profundus, and umbripennis
groups by possessing a distinct diastema
on the masticatory margin of mandible in
the workers and queens, with the excep-
tion of P. trachylissus; but in the workers
of P. trachylissus, five teeth are present on
the masticatory margin, and the pronotum
is armed with a pair of well-developed,
long spines, which are not seen in the
levigatus, profundus, and umbripennis
groups.
The evolution of mandibular dentition
in the workers and queens is one of the
reasons of the diversity of the quadridens
group. The dental formula “apical tooth +
a preapical + a long diastema + two small
basal teeth of similar size”, possessed by
many species of the group, is probably an
“ancestral character”. It has given rise to
three apomorphic dental formulae in the
group: (1) an apical + a preapical + an
intercalary small tooth + two small teeth
of similar size, as in P. trachylissus; (2) an
apical + a preapical + a diastema + three
small teeth of similar size, as in P. eduardi,
P. occultus, and P. quindentatus; and (3) an
apical + a preapical + a long diastema +
a basal tooth (which evolved through the
42]
fusion of the two small teeth), as in P. er-
ythropygus, P. quadridentatus, and P.
wheeleri.
The pronotal armaments in the workers
of this group seem to show an evolutionary
tendency to increase in size. For example,
in P. brevispinosus, the pronotum is armed
with teeth shorter than or about as long as
propodeal armaments. In P. costatus, the
pronotum is armed with a pair of short
spines longer than the propodeal arma-
ments, but much shorter than the distance
between the bases of two pronotal spines.
In P. bicolor, the pronotal spines are very
long, much longer than the propodeal ar-
maments and usually much longer than
the distance between the bases of two pro-
notal spines.
In the workers and queens of the quad-
ridens group, foveolate punctures on the
dorsal head between the frontal carinae
show continuous variation, from a com-
plete absence to several punctures present
to foveolate-reticulate sculpture (or dense
assemblages similar to alveolate sculpture)
or rugoreticulum. For example, P. collinus
and P. flatus workers almost completely
lack foveolate punctures on the dorsum of
head between the frontal carinae. Pristo-
myrmex quadridens workers have scat-
tered foveolate punctures on the dorsum
of head between the frontal carinae, but
the spaces between foveolae are smooth.
Pristomyrmex brevispinosus workers have
rugoreticulum or foveolate-reticulate
sculpture. Some populations, only with fo-
veolate-reticulate sculpture behind the
eyes, are considered intermediate forms
and are grouped into P. brevispinosus.
The males of eight species of this group
are known (Figs. 261-266, 270-276).
These males are more similar to those of
the both trispinosus and levigatus groups
than to the male of P. punctatus (punctatus
group) in palp formula, propodeal arma-
ments, and the shape of petiole or to those
of the wmbripennis group in the size, pro-
podeal armaments, and the shape and
sculpture of petiole.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 42.
Pristomyrmex africanus Karavaiev
Figures 102-108
Pristomyrmex africanus Karavaiev, 1931: 47. Holo-
type worker, Kenya: Mabira (Dogiel) (UENC) [ex-
amined].
Hylidris myersi Weber, 1941: 190. Syntype workers,
Sudan: Equatoria, Aloma Plateau, Khor Aba, 3,700
ft, Lat. 3°47'N/Long. 30°37'E (N. A. Weber) [one
syntype (MCZC) examined]. [Synonymy by Bolton,
1981}.
Pieris myersi subsp. mbomu Weber, 1952: 19. Ho-
lotype worker, Central African Republic: Ubangi-
shari, Bas Mbomu, 5 mi W of Bangassau (N. A.
Weber) (AMNH). [Synonymy by Bolton, 1981].
Hylidris myersi subsp. primus Weber, 1952: 19. Ho-
lotype worker, Zaire: Stanleyville (N. A. Weber)
(AMNH). [Synonymy by Bolton, 1981].
Hylidris myersi subsp. beni Weber, 1952: 20. Syntype
workers, Zaire: 15 mi N of Beni (N. A. Weber) [one
syntype (MCZC) examined]. [Synonymy by Bolton,
1981].
Diagnosis (Worker). Masticatory margin
of Pemdible with a long diastema after the
preapical tooth; ventral surface of clypeus
with two toothlike prominences; eyes con-
taining four to five ommatidia in the lon-
gest row; dorsal surfaces of head between
the frontal carinae and alitrunk with scat-
tered foveolate punctures; petiole and
postpetiole without hairs.
Worker. TL 2.76—3.40, HL 0.74—0.90,
HW 0.74—0.93, CI 98—104, SL 0.68—0.83,
SI 86-95, EL 0.08—0.12, PW 0.48-0.58,
AL 0.68—0.85, PPW 0.22-—0.27, PPL 0.18—
Oe Rel BO=135) 7) = 27):
Mandibles with a few longitudinal basal
rugae. Masticatory margin of mandible
with an apical tooth + a preapical tooth +
a long diastema + a broad and truncated
(or somewhat midconcave) basal tooth.
Basal margin of mandible almost straight,
lacking a foothlike prominence or curved
lobe. Clypeus possessing or lacking a me-
dian longitudinal carina. Anterior clypeal
margin with a median denticle and two
others on each side, but two lateral den-
<
is)
ticles are usually fused into one promi-
nence. Ventral surface of clypeus with two
toothlike prominences. Palp formula 1,3.
Frontal carinae extending to the level of
the posterior margins of eyes. Antennal
scrobes indistinct or very weak. Frontal
lobes absent; thus, the antennal articula-
tions are entirely exposed. Antennal
scapes, when lying on the dorsal head,
close to, or slightly surpassing, the occipi-
tal margin of head. Eyes containing four
to five Gruman in the longest row. Pro-
mesonotum, in dorsal view, sometimes
slightly concave. Profile shape of alitrunk
and pedicel segments as in Figures 103—
104. Pronotum Fusually with a pair of teeth
or short spines but rarely with a pair of
tubercles. Propodeum with a pair of teeth
or short spines. Metapleural lobes round-
ed. Petiole node high in profile, higher
than long, with a long anterior peduncle;
its anterodorsal angle distinctly higher
than the posterodorsal one that is usually
rounded. Subpetiole with a narrow flange.
Postpetiole in profile higher than long,
rounded dorsally, in dorsal view broaden-
ing from front to back and broader than
long. Dorsum of head between the frontal
carinae and the sides of head, with nu-
merous scattered foveolate punctures;
sometimes the punctures are shallow and
sparse. Dorsum of alitrunk smooth, except
for a few foveolate punctures shallow or
conspicuous, present on each side of the
mesonotum. Petiole, postpetiole, and gas-
ter smooth and shining. Dorsum of ead
between the frontal carinae, with some
erect or suberect hairs. Promesonotum
with a few pairs of hairs. Propodeum, dor-
sal surfaces of petiole and postpetiole, and
first gaster tergite lacking erect or suberect
Thatins, A few pairs of forward. -projecting
hairs present on the anterior clypeal mar-
Figures 102-108. Pristomyrmex africanus Karavaiev. 102: Worker head, full-face view; 103: Worker, lateral view; 104: Showing
that pronotal prominences are very weak in some specimens; 105A: Ergatoid queen, full-face view; 105B: Showing two small
denticles on the ventral clypeus; 106: Ergatoid queen, lateral view; 107A: Queen head, full-face view; 107B: Showing two small
denticles on the ventral clypeus; 108: Queen, lateral view.
424
gin. Scapes and tibiae with decumbent
hairs. Color reddish-brown, but gaster
darker.
Ergatoid Queen. TL 3.70, HL 0.94, HW
1.00, CI 106, SL 0.85, SI 85, EL 0.16, PW
0.60, AL 0.92, PPW 0.30, PPL 0.20, PPI
LEO Go = IL)
General shape as in Figures 105—106.
Similar to worker; color and pilosity as in
worker. Sculpture, except for mesonotum
where a few rugae present, as in worker.
The head with three ocelli; eyes larger,
with six to seven ommatidia in the longest
row; pronotum armed with a pair of acute
minute spines; propodeum with a pair of
short spines; mesonotum more convex,
and metanotal groove present. Flight
sclerites and wings lacking, but a black
speck is present on the each lateral margin
of mesonotum.
Queen. TL 3.40-3.82, HL 0.84—0.96,
HW 0.86—1.02, CI 102—107, SL 0.74—0.88,
SI (82-89) EL, 0202021, PW 07/0=0.74;
AL 0.90—1.04, PPW 0.28-—0.30, PPL 0.20—
O22 RPIMal3s2=14 35 — <>)
General shape as in Figures 107—108,
with normal caste differences from the
conspecific worker. Eyes larger. Pronotum
lacking teeth or spines but sometimes with
a pair of blunt tubercles. Mesonotum with
more hairs than in worker. Other charac-
ters similar to worker.
Male. Unknown.
Comments. This African species is close-
ly related to the Asian P. quadridens. Both
worker and queen of P. africanus differ
from those of P. quadridens in having two
teeth on the ventral surface of clypeus and
lacking erect or suberect hairs on both
petiole and postpetiole. The separation of
P. africanus, together with P. trogor, from
the other two Oriental species, P. flatus
and P. collinus, is provided under P. flatus.
The workers of P. africanus and P. trogor
differ from the workers of three Australian
species, P. wheeleri, P. erythropygus, and
P. quadridentatus as follows:
P. africanus and P. trogor
Dorsum and sides of both petiole and
postpetiole lacking erect or suberect
hairs
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Metapleural lobes rounded
Dorsal alitrunk lacking a transverse
ridge at the position of metanotal
groove
P. wheeleri, P. erythropygus, and
P. quadridentatus
Dorsum and sides of both petiole and
postpetiole with erect or suberect
hairs
Metapleural lobes elongate-triangular,
with an apex
Dorsal alitrunk with a transverse ridge
at the approximate position of matan-
otal groove
In addition, in P. africanus and P. trogor
the denticles of the anterior clypeal margin
are smaller but larger and stronger in P.
wheeleri, P. erythropygus, and P. quadri-
dentatus. Pristomyrmex africanus and P.
trogor possess two toothlike prominences
on the ventral clypeus that are not seen in
P. wheeleri and P. erythropygus. Pristo-
myrmex erythropygus has several short ru-
gae on the juncture between the prono-
tum and the mesonotum and has erect or
suberect hairs on the first gastral tergite
that are absent in P. africanus and P. tro-
gor. Pristomyrmex trogor lacks a longitu-
dinal median carina on the clypeus, pos-
sessed by P. wheeleri, P. erythropy
P. quadridentatus.
The workers of P. africanus can be sep-
arated from those of the other four African
Pristomyrmex species as follows: (1) P. af-
ricanus possesses numerous foveolate
o
5
punctures on the dorsal head between the |
frontal carinae that are not seen in P. tro- |
gor and P. orbiceps; (2) P. africanus has |
eyes containing four to five ommatidia in |
the longest row, as compared with eight or
more usually present in P. orbiceps, P. fos-
sulatus, and P. cribrarius; (3) P. africanus
has two teeth on the ventral clypeus that
are absent in P. orbiceps and P. fossulatus; ©
(4) P. africanus lacks coarse longitudinal ©
rugae and erect or suberect hairs on the
petiole and the postpetiole that are pres-
ent in P. cribrarius.
us, and —
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
Distribution. Ghana, Cameroon, Gabon,
Kenya, Zaire, and Angola (Bolton, 1981).
Some more records, including some eco-
logical information, added here (MCZC,
ANIC): Zaire: Ituri F. Beni-Irumu (N. A.
Weber). Sudan: Equatoria, Imatong Mts.
(N. A. Weber). Angola: Falls R. Chicapa,
Saurimo, 9.39°S, 20.24°E, gallery forest,
berlesate (Luna de Carvalho); R. Kahingo,
7.39°S, 20.51°E, gallery forest, berlesate
(Mwaoka); Dundo, R. Mussungue, gallery
for., berlesate (Luna de Carvalho); Dundo,
dry forest (Luna de Carvalho); Dundo,
Carrisso Park, R. Luachimo, 7.22°S,
20.50°E, gallery forest, berlesate (Luna de
Carvalho); R. Camudembele, gallery for-
est, berlesate (Luna de Carvalho); Ghana:
E.R., Mt. Atewa, rainforest, Berlesate (R.
W. Taylor); E.R., Nkwanda For., near En-
yiresi, rainforest, Berlesate (R. W. Taylor);
Tafo, Eastern Reg., rainforest, Berlesate
(R. W. Taylor).
Ecological Information. Weber (1941:
192; 1952: 18-20) noted that “workers
were in rainforest of a luxuriant type re-
ferred to as gallery forest”; they were
among the leaf and humus cover on the
forest floor and were slow moving in habit;
“when disturbed they became motionless,
‘feigning death’ momentarily”.
Pristomyrmex bicolor Emery stat. n.
Figures 109-112
Pristomyrmex trachylissa var. bicolor Emery, 1900:
678. Syntype workers, Sumatra: Si-Rambé,
xii. 1890-iii. 1891 (E. Modigliani) (MCSN, NHMV,
USNM) [examined].
Pristomyrmex taurus Stitz, 1925: 120. Holotype
worker, Philippines: N. Palawan, Binaluan, xi.—
xii.1913 (G. Boettcher) (MNHU) [examined].
Syn. n.
Note: The unique holotype of P. taurus
differs from the syntypes of P. bicolor. In
the former, the pronotal spine length is
0.58, and the anterior half of the dorsum
of the alitrunk is smooth and shining. In
the latter, the pronotal spine length is 0.36
to 0.40, and sometimes only a rather nar-
row smooth area is present between the
bases of two pronotal spines. Pristomyr-
425
mex taurus, however, is here regarded as
a junior synonym of P. bicolor because, af-
ter examining all the specimens available,
I find that the pronotal spine length is con-
tinuously variable in the range of 0.36 to
0.66, and sculpture on the anterior half of
the dorsal alitrunk also shows continuous
variation. I cannot use any lines to separate
this composite at the present. The size of
the holotype of P. taurus is HL 1.10, HW
OS. Cl 9S. SE 114) ST 06) E0220" PV.
O74 AIS) PSE 02 4s Mihensizessor
three syntype workers of P. bicolor are HL
1 20=1228) IW 12421325 Cil03—lO7s sie
1.38—1.46, SI 108-111, EL 0.24—0.26, PW
0.84—0.90, AL 1.40—1.44, PSL2 0.10-0.12.
Diagnosis (Worker). Masticatory margin
of mandible with four teeth and a long di-
astema after the preapical tooth; pronotal
spines exceptionally long (usually 0.40—
0.66), much longer than propodeal arma-
ments (teeth or short spines); dorsum of
head sculptured with coarse rugoreticu-
lum.
Worker. TL 4.58-6.14, HL 1.04—1.42,
HW 1.08-—1.46, CI 98—109, SL 1.14—1.58,
SI 106-120, EL 0.20—0.26, PW 0.65—0.96,
AL 1.18—1.64, PPW 0.33—0.40, PPL 0.38—
0.48, PPI 77-93 (n = 83).
Mandibles usually with several longitu-
dinal rugae, varying from superficial to
rather coarse. Masticatory margin of man-
dible with four teeth arranged as: two ad-
jacent strong apical teeth + a long diaste-
ma + two small basal teeth that are rough-
ly the same size. Basal margin of mandible
lacking a distinctly curved lobe or tooth.
Clypeus with a strong median longitudinal
carina. Anterior clypeal margin sometimes
with seven denticles: a median denticle
and three others on each side, but some-
times one or a few are weak or indistinct,
or two to three of the lateral denticles are
fused into a larger one. Ventral center of
clypeus usually with a transverse ruga of
varying length, sometimes with a very
weak prominence at middle and some-
times without either of these characters.
Palp formula 1,3. Frontal carinae strong,
426
CY SN
RTS
WZ
S40
1)
SE
~~
VO
(cS
Ne
AO
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
SN
Figures 109-112. Pristomyrmex bicolor Emery. 109A: Worker head, full-face view; 109B: Showing a weak transverse ruga on
the ventral clypeus; 110: Worker, lateral view; 111A: Queen head, full-face view; 111B: Showing a transverse ruga on the ventral
clypeus; 112: Queen, lateral view.
extending to the level of the posterior mar-
gins of eyes. Slightly concave scrobal areas
present. Frontal lobes absent; thus, the an-
tennal articulations are entirely exposed.
Antennal scapes, when lying on the dorsal
head, usually surpassing the occipital mar-
gin by one-third to one-fourth of their
length. Eyes usually containing 10 to 12
ommatidia in the longest row. Profile
shape of alitrunk and pedicel segments as
in Figure 110. Pronotal spines well devel-
oped and exceptionally long, varying in
length, usually 0.40 to 0.66, but in a few
specimens they are 0.37 to 0.39. Propo-
deum usually with a pair of acute short
spines, which are 0.12 to 0.20, much more
slender and much shorter than the pro-
notal ones. Propodeal armaments occa-
sionally reduced to a pair of teeth. Both
pronotal and propodeal spines upward —
pointed. Metapleural lobes subtriangular.
In profile, petiole node high, with a long |
anterior peduncle; its anterodorsal angle |
elevated above the posterodorsal. In dorsal |
view, crest of petiole node rounded. Post- !
petiole in profile convex dorsally, in dorsal |
view longer than broad and broadening |
from front to back. Dorsum of head with |
well-developed coarse rugoreticulum. |
Dorsal surface of alitrunk variably sculp- —
tured: At one extreme, in a series from |
North Borneo (SE, Forest Camp, 9.8 km |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 42
SW of Tenom), the dorsum of alitrunk is
entirely rugoreticulate. In a few other se-
ries, the rugoreticulum is absent between
the bases On two pronotal spines, and some
weak rugae or a smooth (broad or narrow)
area are present there. At the other ex-
treme, the sculpture is completely absent,
and the area is smooth and shining on the
anterior half of the dorsum, but the rest of
the dorsal alitrunk is coarsely rugulose.
Sides of pronotum with a rugoreticulum or
many large coarse foveolate punctures.
Sides of the rest of alitrunk irregularly
coarsely rugulose. Dorsal surfaces of peti-
ole node Pal postpetiole smooth and shin-
ing, but a longitudinal ruga present on
Gack side of petiole. Gaster unsculptured,
smooth, and shining. Dorsal surfaces of
head and alitrunk Sei numerous erect or
suberect hairs. Petiole node with a few
(usually two to three) pairs of hairs dor-
sally. Dorsum of postpetiole with at least a
pair of hairs. First gastral tergite lacking
erect or suberect Hears: A row ae forward:
projecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect or suberect hairs. Color usually
reddish-brown but sometimes black-
brown.
Queen. TL 6.54, 6.56; HL 1.34, 1.44;
HW 1.38, 1.50; CI 103, 104; SL 1.48, 1.51;
SI 99, 109; EL 0.32, 0.34; PW 1.20, 1.24:
MG ei 2S 2 RP (OrA1e 0:43. PEEO0:438)
0.48; PPI 85, 90 (n = 2).
General shape as in Figures 111-112,
with normal caste differences from the
conspecific worker; pronotum unarmed.
Other characters similar to worker.
Male. Unknown.
Comments and Discussion. Pristomyr-
mex bicolor occurs in the Oriental region.
It possesses a pair of exceptionally long
pronotal spines, which implies that this
species may have evolved from the ances-
tor of P. costatus. Only slight differences
separate the workers of P. bicolor and of
P. costatus, as follows:
P. bicolor
Pronotal spines well developed and ex-
ceptionally long, usually 0.40 to 0.66
1
(rarely 0.37—0.39), and usually longer
than the distance between their bases
Dorsal surfaces of petiole node and
postpetiole usually with one to three
pairs of hairs
Area between the bases of the pronotal
spines smooth or sculptured; if sculp-
tured, pronotal spine length is over
0.40
Larger species (HW usually 1.20—1.46,
rarely 1.04—1.19; HL usually 1F20=
EAA rarely 1.08—1.19)
P. costatus
Pronotal spines moderately long, usually
0.18 to 0.27 (rarely 0.32), and shorter
than the distance between their bases
Dorsal surfaces of petiole node and
postpetiole with five or more pairs of
hairs
Entire dorsum of alitrunk with devel-
oped rugoreticulum
Smaller species (HW 0.90-1.16, HL
0.91—1.16)
Another alternative is that P. bicolor
may be derived from a P. curvulus—like an-
cestor. Characters separating P. bicolor
from P. curvulus are provided under P.
curvulus.
Pristomyrmex bicolor is also very similar
in appearance to P. trachylissus. The two
species are all from the Oriental region.
The separation of the two species is sum-
marized under P. trachylissus.
It must be pointed out that P. bicolor is
a highly variable species. I cannot separate
any more sibling species from this mass at
the present time. Further collecting and
study will help clarify the situation.
Material Examined (ANIC, BMHH,
BMNH, LACM, MCZC, NHMV). Indo-
nesia: W. Java, 9 km W Djasinga, Dungus
Iwul, lowland rainforest (W. L. Brown):
Java, Bali I, Tjanoi Kuning (J. Winkler);
Borneo: Kaknntan Timur, ITCI Timber
Camp, via Balikpapan, on fallen trees (N.
Johnson); SE Borneo, 17 to 46 km W Ba-
tulitjin, rainforest (W. L. Brown). Malaysia:
North Borneo (SE), Forest Camp, 9.8 km
SW of Tenom (Y. Hirashima); Sabah, Batu
428
Punggul Resort, primary forest, sifting (?);
Sarawak, Genting Highlands (B. Bolton);
Sarawak, 4th Div., G. Mulu Nat. Pk., RGS
Expd., Long pala, lowland rainforest, on
fallen tree and on rotten log (B. Bolton; P.
M. Hammond and J. E. Marshall); Sara-
wak, Mt. Penrissen, 4,500 ft (E. Mjdberg);
Sarawak, Mt. Poi (E. Mjéberg); Malaya,
Sq. Patani (G. H. Lowe); Selangor, Ulu
Gombak For. Reserve, hill forest, Tree
lookout area, ca. 450 m (R. Crozier); Se-
langor, Genting Highlands, below Sri Lay-
an, 900 m, hill forest (W. L. Brown); Neg.
Sembilan, Pasoh For. Res. (M. Brendell,
K. Jackson, and S. Lewis). Philippines: Lu-
zon, Lagunas, Mt. Banahaw above Kina-
buhayan, 600 to 700 m (J. Kodada and B.
Rigova).? : Tjibodas, 1,500 m (?).
Ecological Information. This species oc-
curs in rainforest and has been collected
on fallen trees and rotten logs.
Pristomyrmex brevispinosus Emery
Figures 113-118, 261, 270
Pristomyrmex brevispinosus Emery, 1887: 451. Syn-
type workers and male, Sumatra: Mt. Singalang,
Luglio, 1878 (O. Beccari) [syntype workers
(MCSN, NMMV, USNM) examined].
*Pristomyrmex yaeyamensis Yamane and Terayama,
1999: 17. Holotype worker, Japan: Okinawa Pref.,
Yaeyama Is., Iriomote-jima, 7.viii.1985 (K. Kino-
mura) (MNHA). Syn. n.
Diagnosis (Worker). Masticatory margin
of mandible with at most four teeth; pro-
notal armaments toothlike, about long as
propodeal armaments that are a pair of tri-
* Note: I have seen photographs of Pristomyrmex
yaeyamensis from Japanese Ants Image Database
(which were placed under the name Pristomyrmex
brevispinosus sulcatus). | propose Prystomyrmex yae-
yamensis as a junior synonym of P. brevispinosus for
the following reason: Although P yaeyamensis pos-
sesses an ergatoid queen caste (Yamane and Teraya-
ma, 1999), this condition, at this moment, is not
enough to separate P. yaeyamensis from P. brevispi-
nosus because the type specimens of P. brevispinosus
contain only workers and male. In other words, it is
not known, at the present, whether P. brevispinosus
possesses a normal queen caste, an ergatoid queen
caste, or both. (In Pristomyrmex, some species, e.g.,
P. wheeleri and P. africanus, contain both castes.)
Thus, further ecological investigation is needed.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
angular teeth or short spines; dorsum of
head, at least behind the level of eyes, with
foveolate-reticulate sculpture or rugoreti-
culum.
Workers DIE 3100-426 5s Hn On 3— 18045
HW 0.68—1.04, CI 93-102, SL 0.64—1.06,
SI 94-108, EL 0.14—0.19, PW 0.48-0.68,
AL 0.76—1.10, PPW 0.21—0.27, PPL 0.18—
0.30, PPI 86-113 (n = 100).
Mandibles generally rather smooth but
sometimes with a few longitudinal rugae.
Dentition of the masticatory margin of
mandible arranged as: the strongest apical
+ the second strongest preapical + a long
diastema + two small teeth of similar size
(or a broad tooth with two points). Basal
margin of mandible almost straight, lack-
ing a distinctly curved lobe or tooth. Clyp-
eus with a median longitudinal carina that
is sometimes interrupted. Anterior clypeal
margin with a median denticle and two to
three others on each side. Ventral center
of clypeus usually with an acute tooth, but
sometimes this tooth somewhat low and
broad. Palp formula 1,3. Frontal carinae
strong, extending to the level of the pos-
terior margins of eyes. Antennal scrobes
indistinct, but in some specimens shallow
scrobal areas present lateral to the frontal
carinae. Frontal lobes very weak; thus, the
antennal articulations are almost entirely
exposed. Antennal scapes, when lying on
the dorsal head, slightly surpassing the oc-
cipital margin of head. Eye usually con-
taining seven to eight ommatidia in the
longest row. Profile of alitrunk and pedicel
segments as in Figures 114 and 116. Pron-
otum armed with a pair of toothlike ar-
maments that vary in length, approximate-
ly from 0.06 to 0.10. Propodeum with a
pair of triangular teeth or short spines
varying from 0.04 to 0.12. Metapleural
lobe subtriangular or with a somewhat
rounded apex. Shape of petiole varying: In
some populations, the anterior face of the
petiole node in profile is almost insepara-
ble from the upper surface of the pedun-
cle (Fig. 114), but in other populations,
the anterior face of the petiole node is dis-
tinct from the upper surface of the pedun-
429
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
Figures 113-118. Pristomyrmex brevispinosus Emery. 113A: Syntype worker head, full-face view; 114: Syntype worker, lateral
view; 115A: Non-type worker head, full-face view; 116: Non-type worker, lateral view, showing some variation in the shape of
petiole node and in the length of pronotal armaments; 117A: Queen head, full-face view; 118: Queen, lateral view; 113B, 115B
and 117B: A tooth present on the ventral center of clypeus of syntype worker, non-type worker, and queen, respectively.
430
cle, as shown in Figure 116. Anterodorsal
angle of petiole node higher than the pos-
terodorsal. Postpetiole in profile convex
dorsally, in dorsal view broadening from
front to back. Density and intensity of ce-
phalic sculpture variable: The dorsum of
head is covered fully with coarse forveo-
late-reticulate sculpture in type series, but
with rugoreticulum in the some other se-
ries and sometimes with foveolate-reticu-
late sculpture only behind the eyes. Dor-
sum of the alitrunk showing similar sculp-
tural variation but usually possessing a few
longitudinal coarse carinae. Petiole, post-
petiole, and gaster smooth and shining.
Dorsal surfaces of head and alitrunk with
numerous erect or suberect hairs. Two
pairs of hairs usually present bilaterally on
the dorsum of petiole node. Usually, a pair,
but sometimes two to three pairs, of hairs
on the dorsum of postpetiole. First gastral
tergite lacking erect or suberect hairs. A
few pairs of forward-projecting hairs pre-
sent near the anterior clypeal margin.
Scapes and tibiae with some erect to sub-
erect short hairs. Color reddish-brown to
blackish-brown.
Queen. TL 3.42-4.81, HL 0.82-1.09,
HW 0.79-1.10, CI 93-106, SL 0.76—1.06,
S93 S018 02020272 PWs0:66—0:39!
AL 0.94—1.30, PPW 0.24—0.32, PPL 0.24—
OBS), IPP GOI (py, = IS).
General shape as in Figures 117-118,
with normal caste differences from the
conspecific worker, pronotum unarmed;
other characters similar to worker.
Ergatoid queen. This caste was reported
(Onoyama, 1976; see also Yamane and Ter-
ayama, 1999).
Male (Figs. 261, 270). Two male speci-
mens, together ath 31 workers and six
queens, Sonnet. a series, collected in In-
donesia (SE Celebes, 1-2 km E of Wolasi,
42 km S. Kendari, ca. 350 m), by W. L.
Brown, and each of the two males was
originally mounted, respectively, with a
worker on the same pin: TL 3.10, 3.36; HL
0.60, 0.62; HW 0.61, 0.62; CI 102, 105; SL
0.21, 0.22; SI 35, 39; HWE 0.77, 0.80; EL
OFS 1 OL32- RW OMS OFS 4 AE ae O Ge alaiG:
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
IRIE Ope, OpAalle IRL, O22), Owe IP }l Gls)
95 (n = 2).
Head, including the eyes, distinctly
broader than long. Clypeus convex, with-
out a median longitudinal carina. Palp for-
mula 1,3. Frontal carinae weak and short,
just reaching the level of the posterior
margins of antennal insertions. Maximum
diameter of the median ocellus 0.10 to
0.11. On the mesonotum, notauli pro-
nounced, forming a Y shape; parapsidal
furrows indistinct. Scuto-scutellar sulcus
with 12 to 13 narrow longitudinal ridges.
Middle and hind tibiae without any spurs.
Propodeum slightly tuberculate, lacking
teeth or spines. Metapleural lobe with a
somewhat rounded apex. Petiole node in
profile low and rounded dorsally, with a
fairly long anterior peduncle. Postpetiole
in pr ofile slow and rounded dorsally and in
dorsal view slightly longer than broad.
Dorsum of head smooth and shining. Ali-
trunk smooth and shining, except for those
marked sutures. Petiole, postpetiole, and
gaster smooth and shining. All dorsal sur-
faces with abundant cieet or suberect
hairs. Scapes and tibiae with numerous
erect or suberect hairs. Body reddish-
brown; funicular segments of antennae
light-yellow and wings slightly infuscated.
In general, the male of P. brevispinosus is
extremely similar to the males of both P.
quadridens and P. sulcatus.
Comments and Discussion. Pristomyr-
mex brevispinosus is the most widely dis-
tributed species in the quadridens group.
So far, it has been found in Sumatra, Cel-
ebes, Malaya, Sarawak, Sabah, Thailand,
the Philippines, Taiwan, and Japan.
It is obvious that P. brevispinosus has
evolved from a P. quadridens—like ances-
tor. Pristomyrmex brevispinosus possesses
densely assembled forveolate punctures
(i.e., foveolate-reticulate sculpture) or ru-
goreticulum on the cephalic dorsum in the
workers and queens, on the dorsal alitrunk
in the workers, and on the sides of the
pronotum in the workers and queens. But
P. quadridens has only scattered foveolate
punctures in these areas.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
Some intermediate forms are present
between P. brevispinosus and P. quadri-
dens, but they may be easily assigned to
either of the two species as follow Those
populations possessing a few coarse lon-
gitudinal carinae or some coarse rugae on
the dorsum of the alitrunk or possessing
foveolate-reticulate sculpture only behind
the eyes should be assigned to P. brevispi-
nosus, but those possessing only foveolate
punctures and lacking coarse longitudinal
carinae or rugae on the dorsal Falitrenk
should be assigned to P. quadridens.
The male of P. brevispinosus is indistin-
guishable at present from that of P. quad-
ridens; this strengthens the conclusion that
these two species are closely related. An-
other close relative of P. brevispinosus is
P. sulcatus; their differentiation is dis-
cussed under P. sulcatus. Characters sep-
arating P. brevispinosus from the two Aus-
tralian species, P. foveolatus and P. thora-
cicus, are provided under P. foveolatus.
Pristomyrmex brevispinosus may be
split into two species in the future. One
species would show that the anterior face
of the petiole node, in profile, is insepa-
rable from the upper surface of its anterior
peduncle, and the other species would
show that the anterior face of the petiole
node, in profile, is distinct from the upper
surface of the peduncle and that a concave
shape is present between the two faces.
However, at this moment, I cannot use this
single line to separate the composite.
Note: Lin and Wu (1998) elevated P.
brevispinosus sulcatus var. formosae Forel
to species rank. As an infrasubspecific tax-
on, “formosae” is not an available name in
the genus, according to the International
Code of Zoological Nomenclature (also
see Bolton, 1995: 365). If the specimens
assigned to “formosae” represent a good
species differing from P. brevispinosus, the
correct pr ocean would be to describe it
as new to science. I have, however, ex-
amined three syntypes of “formosae”, and
they fit comfortably with P. brevispinosus
as defined in this revision.
Material Examined (ANIC, BMHH,
431
BMNH, LACM, MCZC, MHNG, NHMYV,
USNM). Indonesia: Sumatra, Pematang,
Siantar (Mann); Si-Rambé (E. Modigliani);
Sulawesi Tengah, near Morowali, Ranu
River Area (M. J. D. Brendell); Sulawesi
Utara, Dumoga-Bone N.P., lowland forest,
200 to 400 m, litter (?); Sulawesi, Dumo-
ga-Bone N.P. (D. F. and A. K. Roche); S
Celebes, Balampesoang Forest, 5 to 8 km
NE Tanete, 400 m, degraded rainforest
(W. L. Brown); SE @llebes. tor 2k
of Wolasi, 42 km S Kendari, ca. 350 m,
rainforest, rotten wood (W. L. Brown); N.
Celebes, Mt. Tangkoko-Batuangus Res. 10
to 200 m, tropical evergreen forest, under
bark log (W. L. Brown); N. Celebes, SW
slope Mt. Klabat, 400 to 600 m, rainforest,
rotten wood (W. L. Brown). Malaysia: Ma-
laya, Genting, Highlands (B. Bolton): Se-
langor, Ulu leona For. Reserve, Univ.
Malaya Field Studies Center, ca. 260 m,
rainforest (R. Crozier); Negri Sembilan,
Sungei Menyala For. Res., near Port Dick-
son, lowland rainforest (W. L. Brown and
Tho Yow Pong); Perak, Sungei, Simei
Falls, Cameron Hlds (T. Jaccoud and P.
Marcuard); Trengganu, Kuala Buka near
Trengganu, berlesate (T. Jaccoud and P.
Marcuard); Sarawak, Semengoh For. Re-
serve, 11 mi SW Kuching, rainforest, nest
ex rotten log (R. W. Taylor); Sarawak, Gn
Matang, 20 km E Kuching, 850 m, sub-
montane forest (L6bl and Burckhardt); N
Borneo (E. Mjoberg); N. Borneo, W. coast
Residency, Ranau, 500 m (T. C. Maa); Sa-
bah, Poring Hot Springs, 500 m (Burck-
har dt and Lobl): Sabah, 7 km N Tambun-
an, 700 m (L6bl and Burckhardt); Sabah,
Batu Punggul Resort, Be forest, itt
ing (?); S Sabah Crocker Range NP., Gg.
eins Highland Res., 1,500 to "1.700 m (>):
Neg. Sembilan Pasoh For. Res. (M. Bren-
dell K. Jackson, and S. Levvis); Pahang,
Genting, Highlands Awana, 1,150 m (Lobl
and Calame); Pahang, Ringlet, ravine,
1,250 m (L6bl and Gulamey Pahang, Batu
Gacs N Kuala, Lumpur (L6bl anid Cala-
me). Philippines: P.R. Mindanao, 2 km N
Malaybalay, 670 m, 8.09°N/125.05°E, re-
forestation area (B. B. Lowery). Thailand:
432
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 119-122. Pristomyrmex collinus sp. n. 119: Worker head, full-face view; 120: Worker, lateral view; 121A: Queen head,
full-face view; 121B: Showing a tooth on the ventral clypeus; 122: Queen, lateral view.
Trang Prov.: Khao Chong Nature Educa-
tion Center, lowland tropical rainforest,
(07.35°N/99.46°E, misc stray foragers, sift-
ed leaf in mixed forest; 07.55°N/99.58°E,
misc stray foragers) (R. R. Snelling and
Saowapa Sonthichari). Taiwan: Pilam (H.
Sauter).
Ecological Information. This species oc-
curs in rainforest and has been collected
on rotten logs.
Pristomyrmex collinus sp. n.
Figures 119-122
Diagnosis (Worker). Pronotum and pro-
podeum each with a pair of short spines;
dorsal surfaces of head and alitrunk un-
sculptured, smooth, and highly polished;
petiole node with one to two pairs of hairs,
and with the anterior face of the node dis-
tinct from the upper surface of peduncle;
HW 0.77-0.94 and HL 0.82-0.94.
Holotype Worker (MCZC). TL. 3.51, HL
0.88, HW 0.84, CI 95, SL 0.86, SI 102,
PW 0.59, AL 0.91. Paratypes, 40 workers
and nine queens (MCZC, BMNH,
MHNG).
Worker. TL 3.36—3.84, HL 0.82—0.94,
HW 0.77-0.94, CI 93-103, SL 0.80—0.94,
SI 98-110, EL 0.14—0.18, PW 0.54—0.62,
AL 0.86-—1.00, PPW 0.25-0.26, PPL 0.25-—
0.28, PPI 89=100 (m= 40).
Mandibles usually smooth and shining
but sometimes with a few longitudinal ru-
gae. Dentition of the masticatory margin
of mandible: the strongest apical + the
second strongest preapical + a long dia-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
stema + two small teeth that are subequal
in size. Basal margin of mandible some-
what straight, lacking a distinct tooth.
Clypeus depressed and smooth, usually
unsculptured but very rarely with a longi-
tudinal median carina. Anterior clypeal
margin usually with a median denticle and
two others on each side but sometimes
with a lateral denticle indistinct or two lat-
eral denticles fused into a larger one. Ven-
tral surface of clypeus with a low, broad-
based, central tooth. Palp formula 1,3.
Frontal carinae extending to the level of
the posterior margins of eyes. Antennal
scrobes absent. Frontal lobes weak; thus,
the antennal articulations are almost en-
tirely exposed. Antennal scapes, when ly-
ing on the dorsal head, slightly surpassing
the occipital margin of head. Eyes mod-
erate, usually containing six to seven om-
matidia in the longest row. Pronotum with
a pair of short but acute spines that are
slightly variable in length. Propodeum
armed with a pair of short spines that are
about equal to or slightly longer than the
pronotal ones. Metapleural lobes subtrian-
gular. In both profile and dorsal view, the
dorsum of alitrunk convex, that is, prono-
tum plus mesonotum forming a convex
dorsum. Petiole node in profile with a fair-
ly long anterior peduncle, its anterodorsal
angle higher than the posterodorsal. Post-
petiole in profile rounded dorsally, in dor-
sal view slightly longer than broad, or
about as long as broad, but always slightly
broadening from front to back. Dorsum of
head, except for a few punctures border-
ing the frontal carinae, smooth and highly
polished. Dorsum of alitrunk unsculptured
and highly polished. Very rarely, the dorsal
surfaces of head and alitrunk with a few
feeble punctures. Petiole, postpetiole, and
gaster smooth and shining. A weak longi-
tudinal ruga usually present on each side
of the petiole but absent in few specimens.
Dorsal surfaces of head and alitrunk with
some sparse erect or suberect hairs. One
or two pairs of hairs present on the dorsal
surfaces of petiole node and postpetiole,
respectively. First gastral tergite lacking
433
hairs. Three pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with erect to
suberect short hairs. Color uniformly red-
dish-brown.
Queen. TL 4.04-4.38, HL 0.87-0.94,
HW 0.85-0.96, CI 94—105, SL 0.84—0.94,
SI 98-102, EL 0.22—0.26, PW 0.76-0.84,
Ale, 0:92=1-268P PW. 0.270 S05 BEE O26=
0.29; PPI 100=111 (m = 8).
Generally similar to worker, except for
normal caste differences. In addition,
pronotum unarmed; propodeal armaments
toothlike, shorter than those of the con-
specific worker.
Male. Unknown.
Comments. This species is known only
from the Philippines so far. It has a num-
ber of relatives. Two of them occur in
Southeast Asia: P. flatus, also from the
Philippines, and P. quadridens, from New
Guinea, Indonesia, and Pohnpei. Of its
more distant five relatives, two, P. african-
us and P. trogor, occur in Africa; and the
other three, P. quadridentatus, P. wheeleri,
and P. erythropygus, are endemic to Aus-
tralia.
The following characters can be used to
separate the workers of P. collinus from
those of P. flatus:
P. collinus
Promesonotum in dorsal view showing a
convex dorsum
Anterior face of petiole node, in profile,
distinct from the upper surface of its
anterior peduncle
Smaller species, with HW 0.77—0.94,
HL 0.82-0.94, EL 0.14—0.18
P. flatus
Promesonotum in dorsal view depressed
or shallow-concave
Anterior face of petiole node not dis-
tinct from the upper surface of the
peduncle
Larger species, with HW 0.98—1.04, HL
0.94—-1.02, EL 0.22-0.24
A list of characters separating P. collinus
and P. flatus from P. quadridens and from
434
TIPE
fis 125
ai B
Figures 123-126. Pristomyrmex costatus sp. n. 123A: Worker head, full-face view; 123B: Showing a transverse ruga on the |
ventral clypeus; 124: Worker, lateral view; 125A: Queen head, full-face view; 125B: Ventral clypeus without a toothlike promi- |
nence; 126: Queen, lateral view.
two African and from three Australian spe-
cies is provided under P. flatus.
Holotype Worker. Philippines: Duma-
guete (J. W. Chapman).
Paratypes. 40 workers and nine queens
with same data as holotype.
Additional Material Examined (BMHH,
USNM, NHMV, MCZC). Philippines: Du-
maguete, Horns of Negros, 3,600 ft (J. W.
Chapman); Dumaguete, Camp (J. W.
Chapman); Los Banos (F. X. Williams):
Luzon, Mt. Makiling (F. X. Williams; L.
Quate and C. Yoshimoto); Luzon, Laguna,
Mt. Makiling, 500 to 1,144 m (H. Zettel);
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Luzon, Laguna, Mt. Banahaw above Kina- |
buhayan, 600 to 700 m (J. Kodada and B. |
Rigova); Romblon Prov., Tablas, S. Agus- |
tin, Dubduban, Busai Falls (H. Zettel); Pa- |
nay Is., forest, 300 m (R. C. Mcqregor). |
Ecological Information. This species oc- |
cur in forest, according to the present re- —
cords.
Pristomyrmex costatus sp. n.
Figures 123-126
Diagnosis (Worker). Ventral surface of —
clypeus lacking a developed tooth, but |
usually with a transverse ruga; pronotal
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
spines fairly long, ca. 1.5 to 2 times the
length of propodeal armaments, but dis-
tinctly shorter than the distance between
the bases of two pronotal spines; dorsal
surfaces of head and alitrunk sculptured
with coarse rugoreticulum; petiole node
lacking foveolate punctures; first gastral
tergite lacking erect or suberect hairs.
Holotype Worker (BMNH). TL 4.14,
HL 0.99, HW 0.96, CI 97, SL 0.96, SI 100,
BE O21 PW. O.6S:7 Al Tl2)) PPW-0:30:
REM Os0O, Per 100: Paratypes, 3 workers
(MCZC): TL 4.48-4.54, HL 1.04—1.08,
ED O:98=1-02) (El 94-945 SI lOve.
SI 108-110, EL 0.20—0.20, PW 0.68-0.72,
Niels Awe ENO S0=Orsle ee EO 33—
0.34, PPI 91-94.
Mandibles usually with several longitu-
dinal rugae. Masticatory margin of man-
dible een four teeth arranged as two ad-
jacent strong apical teeth ie a long diaste-
ma + two emia basal teeth of ernie size.
Basal margin of mandible lacking a dis-
tinctly caved lobe or tooth. Clypeus with
a strong median longitudinal carina. An-
terior clypeal margin ath a median den-
ticle and two to three others on each side.
Ventral surface of clypeus possessing or
lacking a transverse ruga, never armed
with a developed, acute tooth. Palp for-
mula 1,3. Frontal carinae strong, extend-
ing to the level of the posterior margins of
eyes. Slightly concave scrobal areas pres-
ent lateral to the frontal carinae. Frontal
lobes weak so that the antennal articula-
tions are almost entirely exposed. Antennal
scapes, when lying on the dorsal head,
slightly surpassing the occipital margin of
head. Eyes containing 9 to 10 ommatidia
in the longest row. Profile shape of alitrunk
and pedicel segments as in Figure 124.
Pronotum ar med with a pair of str ong and
fairly long spines that are ca. 0.19 to 0. Die
1.5 to 2 enee the length of propodeal ar-
maments, but distinctly shorter than the
distance between the bases of two prono-
tal spines. Propodeum with a pair of acute
short spines that are ca. 0.10 to 0.16 and
more slender than the pronotal ones. Me-
tapleural lobes subtriangular. Petiole node
435
in profile, slightly higher than long, with a
fairly long anterior ‘peduncle: is. antero-
dorsal angle is on a higher level than the
posterdorsal. Postpetiole in profile round-
ed dorsally, in dorsal view broadening from
front to back. Dorsum of head, except for
the scrobal areas where there are only
some transverse rugae, with well-devel-
oped coarse rugoreticulum. Similar sculp-
ture present on the sides of pronotum and
the dorsum of alitrunk. Petiole with a
coarse longitudinal ruga on each side, but
dorsum es petiole mode unsculptured and
smooth. Dorsum of postpetiole unsculp-
tured and smooth. Gaster smooth and
shining. Dorsal surfaces of head and ali-
trunk with numerous erect or suberect
long hairs. Sides and dorsum of petiole
node and postpetiole with five or more
pairs of hairs in the type specimens. First
gastral tergite lacking erect or suberect
hairs. Several pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with some erect
to suberect hairs. Color reddish-brown.
Queen. Two queens, respectively, from
N. Borneo and Sarawak, with the following
measurements: TL 5.24, 5.26: HL 1. 12,
202 FW, Wil6: 122 Cl l02) L042 SIE es:
1.20: SI 98, 102: EL 0.28, 0.31: PW 1.00;
NO AeA O2 SP PNWViOrsS 30:44 s Pils
0.40, 0.44, PPI 95, 100.
General shape as in Figures 125-126,
with normal caste differences from the
worker, pronotum unarmed; other char-
acters similar to worker.
Male. Unknown.
Comments and Discussion. Pristomyr-
mex costatus is extremely similar in ap-
pearance to the another Oriental species,
P. sulcatus. The workers and queens of the
two species can be separated as follows: In
P. costatus, the ventral surface of the clyp-
eus lacks a toothlike prominence, usually
with a transverse ruga; in P. sulcatus, the
ventral center of the clypeus has an acutely
prominent tooth and no transverse ruga.
The separation of P. costatus from P.
modestus and from P. bicolor is summa-
rized under the latter names.
436 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
@ oP. costatus
* Ps SulleeEeus
Figure 127. Distributions of Pristomyrmex costatus and Pristomyrmex sulcatus.
Whether or not P. costatus and P. sul- Borneo, and the Philippines but P. sulcatus
catus have a relationship of allopatric dis- in Pahang of Malaya, Thailand, Nepal,
tribution needs further studies. According Burma, and China. Further, whether P.
to the present records, as shown in Figure sulcatus, P. costatus, and P. brevispinosus
127, P. costatus occurs in Negri Sembilan evolved from the results of character dis-
of Malaya, Singapore, Sabah, Sarawak, placement also needs further studies (P.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
costatus is more distant to sympatric IP,
brevispinosus than to P. sulcatus).
The following additional material shows
some interesting variation: In nine work-
ers, the dorsums of petiole node and post-
petiole have five or more pairs of hairs, but
the ventral surface of clypeus has only a
short ruga. Four workers from Singapore
have two pairs of hairs on the dorsum of
petiole node and a pair on the dorsum of
postpetiole. In a worker from Sabah, the
anterodorsal angle of the petiole node in
profile is not distinctly higher than the
posterodorsal. In three workers, one from
Sarawak and the other two from Sabah,
the ventral center of clypeus lacks any
transverse ruga but possesses a minute
prominence instead of an acute tooth, and
the dorsal surfaces of petiole and postpe-
tiole have more than seven pairs of hairs.
A single specimen, from the Philippines,
shows reduced sculpture of the dorsal sur-
faces of head and alitrunk. The measure-
ments of the previously described speci-
mens are TL 3.94-4.54, HL 0.91-1.18,
HW 0.90-1.13, CI S9—103, SL 0.96—1.21,
SI 100-112, EL 0.18—0.24, PW 0.60-0.76,
AL 0.98—1.30 (n = 18).
In addition, a series from Borneo ap-
pears to be intermediate in size between
P. costatus and P. bicolor by possessing the
following worker measurements: TL 4.64—
Se, IRE IOO SIZE JEW OR See Olt
931045 7S a4 22 SI 1038=113. EE
0:20-0:22, PW 0.70=0.78, AL 1.16—1.37,
PSbLil 0:28-0:32, PSL2 0.10=0.116 (7 = 25).
It is possible that the previously de-
scribed material may comprise one or
more sibling species. Further collecting
and biological investigation are needed to
resolve this possibility.
Holotype Worker. Malaysia: Neg. Sem-
bilan, Pasoh For. Res., xi.1994, (M. Bren-
dell, K. Jackson, and S. Lewis).
Paratypes. Three workers, N. Borneo
(E. Mjéberg).
Records of the Previously Examined
Non-Type Material (ANIC, BMHH,
BMNH, MCZC, MHNG, NHMV). Singa-
pore: Nee Soon, Swamp forest, rainforest,
437
nest ex rotten log (R. W. Taylor); Bukit Ti-
mah Nat. Res., degraded coastal hill forest,
on granite (D. H. Murphy). SE Borneo:
17 to 46 km W Batulitjin, lowland rainfo-
rest (W. L. Brown); Borneo: Pajan (E.
Mjéberg). Sarawak: Gunong Matang 120
m (T. C. Maa); 4th Div., G. Mulu Nat. Pk.,
RGS Expd., Long pala, lowland rainforest,
on rotten log and in leaf litter (B. Bolton);
Semengoh NSG, 30 km S Kuching (H.
Zettel). Sabah: Batu Punggul Resort pri-
mary forest, sifting (P); 43 mi, labuk Rd.
ex Sandakan (Lungmanis) (R. W. Taylor);
7 km N Tambunan, 700 m (L6bl and
Burchhardt); N. Borneo, Tutu River (E.
Mjéberg); N. Borneo, (SE) Forest Camp,
9.8 km SW of Tenom (K. J. Kuncheria).
Philippines: Mindanao, Davao Province,
Mt. McKinley, E. slope, 3,300 ft, under
bark (F. G. Werner).
Note: An unusual worker (BMNH), col-
lected from Sabah (K. K.-Tambunan,
Crocker Range, 1,600 m), having a smooth
patch present between the bases of two
shorter pronotal spines, is tentatively
placed under P. costatus; the size of the
specimen is TL 3.84, HL 0.96, HW 0.92,
CI 96, SL 0.96, SI 104, EL 0.18, PW 0.64,
AL 0.96, PSLI1 ca. 0.16, PSL2 ca. 0.08.
Ecological Information. See the section
“Records of the Previously Examined
Non-Type Material”.
Pristomyrmex curvulus sp. n.
Figures 128-131
Diagnosis (Worker). Pronotal spines ex-
ceptionally long, about 0.38 to 0.44; pro-
podeal spine length ca. 0.12 to 0.16; an-
terior face of petiole node distinctly sepa-
rable from the upper surface of peduncle;
dorsal surfaces of head and _ alitrunk
smooth and shining.
Holotype Worker (MCZC). TL5.24, HL
eis; AW 1.20; Cl 102, Sk. 1.36, SP Hs:
EL 0.22, PW 0.79, AL 1.26. Paratypes, 34
workers and one queen (MCZC, BMNH,
LACM, MHNG).
Worker. TL 4.62-5.30, HL 1.08—1.26,
HW, 1_08=1-25,, Cl 97-105; SL, 122-4
SI 106-117, EL 0.20—0.26, PW 0.74—0.82,
438
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 128-131.
Pristomyrmex curvulus sp. n. 128A: Worker head, full-face view; 128B: Showing a transverse ruga on the
ventral clypeus; 129: Worker, lateral view; 130A: Queen head, full-face view; 130B: Showing a transverse ruga on the ventral
clypeus; 131: Queen, lateral view.
Mlle tO Me PNVOS0=0.34 Pel, Oloa—
0.40, PPI 78-94 (n = 20).
Mandibles generally smooth and _ shin-
ing, with a few basal longitudinal rugae.
Masticatory margin of mandible with four
teeth: an apical + a preapical + a long
diastema + two small denticles that are
roughly the same size. Basal margin of
mandible lacking a toothlike prominence.
Clypeus shining, with a median longitudi-
nal carina; sometimes a few additional su-
perficial rugae present. Ventral surface of
clypeus usually with a long transverse ruga.
Anterior clypeal margin usually with seven
denticles (a median one and three others
on each side), but in some specimens, one
to two denticles weak or rudimentary. Palp
formula 1,3. Frontal carinae just extending
to the level of the posterior margins of
eyes. Slightly concave scrobal areas pres-
ent lateral to the frontal carinae. Frontal
lobes weak so that the antennal articula-
tions are almost entirely exposed. Antennal
scapes long, surpassing the occipital mar-
gin by one-fourth to one-third of their
length. Eyes usually containing over 10
ommatidia in the longest row. Profile
shape of alitrunk and pedicel segments as
in Figure 129. Pronotum armed with a
pair of exceptionally long spines that are
about 0.38 to 0.44 and longer than the dis-
tance between their bases. Propodeum
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
with a pair of acute short spines that are
ca. 0.12 to 0.16, about as long as the dis-
tance between their bases, and shorter
than 0.5 times pronotal spine length. Both
pronotal and propodeal spines directed
upward. Metapleural lobes subtriangular.
Petiole in profile nodiform, with a long an-
terior peduncle; the anterior face of the
node distinctly separable from the upper
surface of its anterior peduncle, and its
dorsum sloping somewhat downward pos-
teriorly. Postpetiole in profile rounded
dorsally, in dorsal view distinctly longer
than broad and broadening from front to
back. Dorsum of head generally smooth
and shining, but gena with a few foveolate
punctures, and frontal area usually with a
few weak short rugae. Dorsal alitrunk,
petiole, and postpetiole unsculptured,
smooth, and shining. Gaster unsculptured.
Dorsal surfaces of head and alitrunk with
numerous erect or suberect hairs. Petiole
node and postpetiole each with some hairs
as shown in Figure 129. Antennal scapes
and tibiae with numerous erect or sub-
erect hairs. First gastral tergite without
erect or suberect ais. A re pairs of for-
ward-projecting hairs present near the an-
terior clypeal margin. Color reddish brown
but light yellow in a few specimens.
Queen. TL 6.16, HL 1.18, HW 1.28, CI
IOS, Sls ILS, SIMO IaIly O28, IPWY ILL
Mle O2) PW 0:40> PPE O44. PPT SING
= Il).
General shape as in Figures 130-131,
with normal caste differences from con-
specific worker; pronotum unarmed; peti-
ole with a lateral longitudinal ruga on each
side. Other characters similar to worker.
Male. Unknown.
Comments and Discussion. This species
is closely related to P. longispinus, also
from Dumaguete, Philippines, but the
workers of Hhieee two species can be sep-
arated by the following characters:
P. curvulus
Anterior face of petiole node, in profile,
distinct from the upper surface of pe-
duncle
439
Clypeus with a median longitudinal ca-
rina
Frontal carinae extending to the poste-
rior margins of eyes
Propodeal spines shorter, about as long
as the distance between their bases.
Anterior clypeal margin usually with
seven small denteles
P. longispinus
Anterior face of petiole node, in profile,
not distinct from the upper surface of
peduncle
Clypeus lacking a median longitudinal
carina
Frontal carinae not extending to the
posterior margins of eyes
Propodeal spines longer, about two to
three times the distance between
their bases in length
Anterior clypeal margin usually with
three to five denticles
Pristomyrmex curvulus may also have a
close relationship with P. bicolor. The
workers and queens of the two species can
be separated as follows: In P. curvulus, the
dorsal surfaces of the head and the ali-
trunk are smooth and shining, with only a
few shallow foveolate punctures present
on the genae and a few short rugae on the
frontal area, whereas in P. bicolor, the dor-
sum of the head is entirely covered with
coarse rugoreticulum, and the dorsum of
the alitrunk is also strongly sculptured with
coarse rugae.
It is possible that P. curvulus is derived
from a P. collinus—like ancestor. The work-
er of P. curvulus, apart from the excep-
tionally long pronotal spines, a transverse
ruga on the ventral surface of the clypeus,
and its larger size, is similar to that of P.
collinus. The queen of P. curvulus is also
similar to that of P. collinus, but in the for-
mer it is larger, and the ventral surface of
the clypeus ahas a transverse ruga, not a
toothlike prominence as in P. cole
Holotype Worker. Philippines: Duma-
guete (J. W. Chapman).
Paratypes. Philippines: two workers,
Dumaguete, 7.vi.1942 (J. W. Chapman);
440
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
ZA
EST S
Figures 132-133. Pristomyrmex eduardi Forel. 132A: Worker head, full-face view; 132B: Showing a toothlike prominence on
the ventral clypeus; 133: Worker, lateral view.
two workers, Dumaguete, 14.vi.1942 (J. W.
Chapman); 12 workers, Dumaguete,
10.v.1947 (J. W. Chapman); two workers,
Dumaguete, 4.v.1948 (J. W. Chapman);
one worker, Dumaguete, 1949 (J. W.
Chapman); two workers and one queen,
Dumaguete, 1950 (J. W. Chapman); 11
workers, Dumaguete, Horns of Negros,
3,600 ft (J. W. Chapman); two workers,
Dumaguete, Horns of Negros, 3,600 ft
(Domingo Empeso).
Additional Material Examined (MCZC).
Some specimens, also collected in Duma-
guete, Philippines, by J. W. Chapman, are
not included in the type series because
they are badly mounted or damaged.
Ecological Information. Unknown.
Pristomyrmex eduardi Forel
Figures 132-133
232
Pristomyrmex eduardi Forel, 1914: 232. Holotype
worker, Sumatra Oriental, Bah Boelian (M. v. But-
tel) (MHNG) [examined].
Diagnosis (Worker). Masticatory margin
of mandible with five teeth; pronotum un-
armed; eyes with three to four ommatidia
in the largest row.
Worker. TL 2.9, HL 0.77, HW 0.74, CI
96, SL 0.68, SI 92, EL 0.07, PW 0.50, AL
0.78) (= 1):
Mandibles smooth and shining, except
for a few longitudinal rugae. Masticatory
margin of mandible with five teeth ar-
ranged as the strongest apical + the sec-
ond strongest preapical + a diastema +
three small denticles of similar size; the
length of diastema is about equal to the
distance covered by three small denticles.
Basal margin of mandible lacking a tooth-
like prominence. Clypeus depressed, with
a short median carina that does not reach
the anterior clypeal margin but runs
through the frontal area. Anterior clypeal
margin with five toothlike prominences;
the median three somewhat truncated.
Ventral center of clypeus with a prominent
tooth. Frontal carinae strong, extending to
the level of the posterior margins of eyes
and forming the dorsal margins of the
shallow scrobes. Frontal lobes weak. Eye
small, with three to four ommatidia in the
longest row. Occipital margin in full-face
view feebly concave. Profile of alitrunk and
pedicel segments as in Figure 133. Pron-
otum unarmed, lacking a pair of teeth or
spines. Propodeum armed with a pair of
acute short spines. Metapleural lobes
prominent and rounded. Petiole in profile
view with a fairly long anterior peduncle;
the anterodorsal angle of the node high,
and its dorsum sloping downward poste-
riorly. Postpetiole in profile with a round-
ed dorsum. Dorsum of head with coarse
rugoreticulum, except for a smooth, me-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 44]
152°25'E, in rotten logs, xi.1957 (Darlingtons)
dian longitudinal strip. Dorsum of ali-
trunk, as well as two sides of pronotum,
with developed rugoreticulum. Petiole and
postpetiole smooth and shining. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with numerous erect or suberect
short hairs. Two pairs of the similar hairs
present on the dorsum of petiolar node
and three pairs on the dorsum of postpe-
tiole as shown in Figure 133. First gastral
tergite lacking erect or suberect hairs. A
few pairs of forward. -projecting hairs pres-
ent near the anterior clypeal margin.
Scapes and tibiae with some erect to sub-
erect short hairs. Color reddish-brown.
Queen and Male. Unknown.
Comments. This species is known only
from the holotype. Its two close relatives
are P. quindentatus, from Indonesia, and
P. occultus, from Indonesia and Malaysia.
Pristomyrmex eduardi can be separated
from P. quindentatus and P. occultus be-
cause it lacks pronotal armaments and pos-
sesses smaller eyes (EL = 0.07, with three
to four ommatidia in the longest row) in
the workers.
Though it was considered by Forel
(1914) to be similar to P. punctatus (=P.
pungens = P. japonicus), P. eduardi cannot
be placed in the punctatus group because
(1) it has five teeth present on the masti-
catory margins of the mandibles, (2) its
eyes are very small, (3) the ventral center
of the clypeus is equipped with a promi-
nent tooth, and (4) the petiole node has a
distinct anterior face. In addition, its pro-
podeal spines are much shorter than those
in the four Oriental species of the punc-
tatus group. Incidentally, the palp formula
of P. eduardi cannot be determined from
the unique holotype and thus remains un-
known at this time.
Ecological Information. Unknown.
Pristomyrmex erythropygus Taylor
Figures 134-137
Pristomyrmex erythropygus Taylor, 1968: 65. Holo-
type worker, Australia: NE. New South Wales, Aca-
cia Biareauh near Old Koreelah, ca. 28°24'S,
(ANIC) [examined].
Diagnosis (Worker). Masticatory margin
of fe aadible with three teeth: anterior
clypeal margin with three strong teeth;
propodeal armaments, ca. 0.13 to 0.20,
usually slightly longer than pronotal
spines; dorsum of head smooth, but dorsal
alitrunk with several longitudinal rugae
present at the juncture between the pron-
otum and the mesonotum; first gastral ter-
gite usually with erect or suberect hairs.
Worker. TL 3.48-3.90, HL 0.90-1.08,
HW 0.94—1.18, CI 104-110, SL 0.92—1.00,
SI 85-97, EL) 0:16-0:20; PW 0:56—0.66;
AL 0.84—0.98, PPW 0.24—0.30, PPL 0.19—
023) BRAS lO — 8):
Mandibles generally smooth and _ shin-
ing, but sometimes with a few basal lon-
gitudinal rugae. Dentition of the mastica-
tory margin rer mandible: an apical tooth
+ a preapical + a long diastema + a some-
what truncated basal tooth. Basal margin
of mandible lacking a distinctly curved
lobe or tooth. Clypeus with a median lon-
gitudinal carina. Anterior clypeal margin
with three teeth: a median denticle and
one on each side. Ventral surface of clyp-
eus with a short transverse carina or with
a low, broad prominence. Palp formula
2,2. 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
entirely exposed. Antennal scapes, laid on
the dorsal head, slightly surpassing the oc-
cipital margin of head. Eyes containing
eight to nine ommatidia in the longest row.
Profile shape of alitrunk and pedicel seg-
ments as in Figure 135A. Pronotum armed
with a pair of moderately long spines, vary-
ing in length from 0.08 to 0.13. Propodeal
spines usually slightly longer than pronotal
ones, varying in length from 0.13 to 0.20.
Metapleural lobes triangular and much
shorter than propodeal spines. Petiole
node in profile with the anterodorsal angle
higher than the posterodorsal. Anterior
and dorsal faces of the postpetiole in pro-
442
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
SY
Ss
NY!
mee
oe
OIA
137
Figures 134-137. Pristomyrmex erythropygus Taylor. 134A: Worker head, full-face view; 134B: Showing a short transverse
carina or a low, broad prominence on the ventral clypeus; 135A: Worker, lateral view; 135B: Dorsum of the worker alitrunk,
dorsal view; 136A: Queen head, full-face view; 136B: Showing a short transverse ruga, or a low, broad prominence on the
ventral clypeus; 137: Queen, lateral view.
file forming a single curved surface; in dor-
sal view, postpetiole distinctly broader
than long. Dorsum of head smooth and
shining, except for a few short rugae pres-
ent below the frontal carinae around the
antennal fossae and on the genae. Dorsum
of alitrunk possessing (1) several short ru-
gae present approximately at the juncture
between the pronotum and the mesono-
tum (but weak in a smaller specimen), (2)
a few transverse rugae present near the
anterior pronotal margin, and (3) a trans-
verse ridge present at the approximate po-
sition of metanotal groove. Petiole, post-
petiole, and gaster smooth and _ shining.
Dorsal surfaces of head and alitrunk with
numerous erect or suberect hairs. Dorsal
surfaces of petiole node and postpetiole,
respectively, with a pair of bilaterally dis-
tributed long hairs; sometimes the crests
of petiole node and postpetiole with ad-
ditional one to two pairs of short hairs.
First gastral tergite with numerous, evenly
distributed, erect or suberect hairs. (Note:
In three specimens placed under P. ery-
thropygus, several longitudinal rugae are
present at the juncture between the pron-
otum and the mesonotum, but erect or
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
443
Figures 138-139. Pristomyrmex flatus sp. n. 138A: Worker head, full-face view; 138B: Showing a very short ruga on the ventral
clypeus; 139: Worker, lateral view.
suberect hairs are absent from the first
gastral tergite. Are these hairs artificially
erased? Further collecting is needed to
clarify this question.) A few pairs of for-
ward-projecting hairs present near the an-
terior clypeal margin. Scapes and _ tibiae
with some erect to suberect short hairs.
Color reddish-brown to blackish-brown.
Queen. TL 4.40, HL 1.14, HW 1.27, CI
SO oly SON Ei nOl225 PW 40:s6:
AL 1.20, PPW 0.34, PPL 0.24, PPI 142 (n
= jl),
General shape as in Figures 136-137,
with normal caste differences from the
conspecific worker; pronotum unarmed;
pro-mesonotum lacking longitudinal ru-
gae; propodeal spines distinctly shorter
than those in conspecific worker; other
characters similar to those in the conspe-
cific worker.
Male. Unknown.
Comments. Pristomyrmex erythropygus
is a sibling species of P. wheeleri and also
related to P. quadridentatus. The three
species are all from Australia. Pristomyr-
mex erythropygus differs from P. wheeleri
and P. quadridentatus because the former
possesses numerous erect or suberect
hairs on the first gastral tergite and several
short longitudinal rugae at the juncture
between the pronotum and the mesono-
tum in the workers that are absent in the
latter two species. In addition, the propo-
deal spines are usually slightly longer than
the pronotal ones in the workers of P. er-
ythropygus but much shorter than the
pronotal spines in P. quadridentatus.
The differences between P. erythropy-
gus and the two Oriental species (P. flatus
Al P. collinus) and between P. erythro-
pygus and the two African species (P. af-
ricanus and P. trogor) are mentioned un-
der P. flatus and P. africanus, respectively.
Material Examined (ANIC, MCZC).
Australia: New South Wales, Acacia Pla-
teau, in rotten logs (Darlingtons); NE New
South Wales, Nothofagus Mt., via Wood-
enbong, Nothofagus forest, 1,100 m,
sieved Titter, Q. M. Berlesale No. 414 (G.
Monteith and G. Thompson); NE New
South Wales, Gibraltar Range N.P., rain-
forest, granite, sieved litter, QM. Berlesate
No. 270) (Gab: Nionterth): New South
Wales, Gibraltar Range Nat. Pk., 920 m,
QO SileS M5222 2h) Beviecat ANIC 836,
closed forest leer (L. Hill); New South
Wales, 10.5 km W of Gibraltar Ra N.P.
(HQ), rainforest, sieved litter, QM. Ber-
lesate No. 213 (G. B. Monteith).
Ecological Information. This species oc-
curs in rainforest and has been collected
in rotten logs and in litter berlesates.
Pristomyrmex flatus sp. n.
Figures 138-139
Diagnosis ( Worker). Pronotum and pro-
podeum each with a pair of short spines;
444
dorsal surfaces of head and alitrunk
smooth and unsculptured; petiole node
with at least two pairs of hairs; anterior
face of petiole node indistinguishable from
the upper surface of its anterior peduncle;
HW 0.98-1.04 and HL 0.94—1.02.
Holotype Worker (MCZC). TL 3.94, HL
1.02, HW 1.04, CI 102, SL 1.10, SI 106,
EL 0.24, PW 0.66, AL 1.03. Paratypes,
three workers (MCZC, BMNB#H).
Worker. TL 3.79-4.14, HL 0.94—1.02,
HW 0.98—1.04, CI 102—106, SL 1.02—1.12,
SI 104-108, EL 0.22—0.24, PW 0.64—0.67,
AL 1.02-1.08, PPW 0.28-—0.30, PPL 0.28—
0.30, PPI 93-100 (n = 4).
Mandibles generally smooth and shin-
ing, with a few basal short rugae. Denti-
tion of the masticatory margin of mandi-
ble: the strongest apical tooth + the sec-
ond strongest preapical + a long diastema
+ two small denticles that are about equal
in size. Basal margin of mandible lacking
a toothlike prominence. Clypeus de-
pressed and smooth, but the frontal area
with a median carina that extends a little
to the clypeus. Anterior clypeal margin
usually with a median denticle and three
other small denticles on each side, but
sometimes one of the lateral denticles very
weak and indistinct. Ventral center of clyp-
eus with a weak, toothlike prominence.
Palp formula 1,3. Frontal carinae approx-
imately reaching to the level of the pos-
terior margins of eyes. Antennal scrobes
absent. Frontal lobes very weak so that the
antennal articulations are almost com-
pletely exposed. Antennal scapes rather
long, when lying on the dorsal head, sur-
passing the occipital margin by one-fifth to
one-fourth of their length. Eyes containing
eight to nine ommatidia in the longest row.
Pronotum armed with a pair of short ro-
bust spines as in Figure 139. Propodeum
with a pair of slender acute spines that are
directed upward and slightly longer than
the pronotal ones. Metapleural lobes sub-
triangular. Pronotum and mesonotum in
dorsal view slightly concave between the
pronotal spines and between the two lat-
eral margins of mesonotum, respectively.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Petiole node massive; its anterior face in
profile indistinguishable from the upper
surface of petiole peduncle (Fig. 139).
Dorsum of petiole node in dorsal view
about rounded. Postpetiole in profile con-
vex dorsally, in dorsal view slightly longer
than broad or about as long as broad,
broadening from front to back. Dorsum of
head usually smooth and shining, but some
small and shallow hair pits present. Dor-
sum of alitrunk unsculptured, smooth and
shining. Sides and dorsum of petiole and
postpetiole smooth and shining. Gaster
unsculptured. Dorsal surfaces of head with
numerous erect or suberect hairs. Dorsum
of alitrunk with sparse erect to suberect
hairs. Two pairs of hairs present on the
dorsum of petiole node, and two to three
pairs on the dorsum of postpetiole. First
gastral tergite lacking erect or suberect
hairs. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with numerous
erect to suberect short hairs. Color red-
dish-brown but sometimes yellow-brown.
Queen. Unknown.
Male. See the following note.
Comments. Like P. collinus, P. flatus is
also known only from the Philippines.
Pristomyrmex flatus is similar to P. colli-
nus, P. quadridens, P. africanus, P. trogor,
P. quadridentatus, P. wheeleri, and P. ery-
thropygus. The workers of P. flatus can be
immediately recognized; because in P. fla-
tus, the anterior face of the petiole node
is not distinct from the upper surface of
its anterior peduncle, which is different in
the previously mentioned species.
A more detailed discussion of the sep-
aration of P. flatus from P. collinus is pro-
vided under P. collinus.
Pristomyrmex flatus and P. collinus dif-
fer from P. quadridens in the workers as
follows:
P. flatus and P. collinus
Dorsum of alitrunk unsculptured,
smooth, and shining |
Dorsum of head smooth and shining, |
|
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
except for a few punctures bordering
the frontal carinae
P. quadridens
Dorsum of alitrunk with scattered fo-
veolate punctures; sometimes some
short rugae present
Dorsum of head with a few to many fo-
veolate punctures, except for those
bordering the frontal carinae
Pristomyrmex flatus and P. collinus dif-
fer from two African species, P. africanus
and P. trogor, in the workers as follows:
P. flatus and P. collinus
Ventral center of clypeus with a short
ruga or small toothlike prominence
At least one to two pairs of hairs present
on the petiole node and postpetiole
Metapleural lobes triangular
P. africanus and P. trogor
Ventral surface of clypeus with two
teeth
Petiole and postpetiole lacking erect or
suberect hairs
Metapleural lobes rounded
The other differences include that the
anterior face of the petiole node is distinct
from the upper surface of the peduncle in
the workers of P. africanus and P. trogor
but almost indistinct in P flatus; the dor-
sum of the alitrunk in dorsal view is convex
in the workers of P. collinus but distinctly
shallowly concave in P. africanus and P.
trogor; the dorsum of the head is sculp-
tured with scattered foveolate punctures in
the workers of P. africanus but is generally
smooth and shining in P. flatus and P. col-
linus; the frontal carinae extend to the lev-
el of the posterior margins of eyes in the
workers of P. flatus and P. collinus but not
so in P. trogor.
Pristomyrmex flatus and P. collinus dif-
fer from three Australian species P. quad-
ridentatus, P. wheeleri, and P. erythropy-
gus in the workers as follows:
P. flatus and P. collinus
Anterior clypeal margin with five to sev-
en smaller denticles
445
Clypeus unsculptured
Masticatory margin of mandible with
four teeth, consisting of an apical, a
preapical, and two small basal denti-
cles
Palp formula 1,3.
Alitrunk in dorsal view unsculptured
P. quadridentatus, P. wheeleri, and
P. erythropygus
Anterior clypeal margin with three
strong and larger teeth
Clypeus sculptured with a strong me-
dian carina
Masticatory margin of mandible with
three teeth, consisting of an apical, a
preapical, and a basal tooth
Palp formula 2,2
Alitrunk in dorsal view with a transverse
ridge present at the approximate po-
sition of metanotal groove
In addition, the anterior face of the pet-
iole node, in profile, is distinct from the
upper surface of the peduncle in the work-
ers of P. quadridentatus, P. wheeleri, and
P. erythropygus but almost indistinct in P.
flatus; the dorsum of the alitrunk is convex
in the workers of P. collinus but almost flat
or shallowly concave in P. quadridentatus,
P. wheeleri, and P. erythropygus.
Holotype Worker. Philippines: Luzon I.,
Bauqui; xi.1923. (R.C.Mcq.).
Paratypes. Three workers with same
data as holotype.
Ecological Information. Unknown.
Note: The following five male speci-
mens, with same data as holotype and
paratypes, may represent the male of this
species. I tentatively place these males un-
der P. flatus, which needs further confir-
mation.
Male (Figs. 262, 271). TL. 3.62—3.94, HL
0.64—0.68, HW 0.56-0.59, CI 85-91, SL
0.22-0.25, SI 39-43, HWE 0.98-1.04, EL
04820) 512 RWG074=OlS0n AealelG=1e 24s
PPW 0.24-0:24" PPL 0.24—0.24 PPI 100=
100 (n = 5).
Head, including the eyes, much broader
than long; while excluding the eyes, dis-
tinctly longer than broad. Eyes very large
446
and prominent; their length is about three-
fourths of the head length. Clypeus con-
vex, somewhat semicircular, its anterior
margin straight and posterior one semicir-
ular Palp for mula 1,3. Frontal carinae ab-
sent or short, slightly beyond the anterior
margins of antennal insertions. F rontal
area usually with a median longitudinal ca-
rina. Ocelli developed; maximum diameter
of median ocellus 0.16 to 0.18. On the me-
soscutum, notauli distinct, forming a Y
shape, but usually without distinct ridges
in them; parapsidal furrows absent. Scuto-
scutellar sulcus usually with 9 to 10 narrow
ridges. Propodeum lacking armaments.
Metapleur: al lobes subtriangular. Middle
and hind tibiae without any spurs. Petiole
node in profile low, with a subtriangular
apex and a rather long anterior peduncle;
anterior face of the node, together with
the dorsal surface of its anterior peduncle,
forming a long declivity, which reaches the
top of fhe nen Postpetiole in profile low,
rounded dorsally, in dorsal view about as
broad as long. Dorsum of head smooth
and shining, except for a few short rugae
present on ‘the posterior margin of clypeus.
Alitrunk generally smooth antl shining, ex-
cept for those marked sutures. Petiole,
postpetiole, and gaster unsculptured,
smooth, and shining. All dorsal surfaces
with abundant long hairs. Scapes and tib-
iae with numerous erect or suberect short
hairs. Color reddish-brown; hairs reddish-
brown; antennae sometimes yellow-brown;
wings somewhat infuscated.
Pristomyrmex foveolatus Taylor
Figures 140-143
Pristomyrmex foveolatus Taylor, 1965: 38. Holotype
worker, Australia: N. Queensland, west of Tully,
Clump Point, rainforest, a few feet above sea level,
25.vi.1962 (R. W. Taylor) (MCZC) [examined].
Diagnosis (Worker). Pronotum with a
pair of triangular short spines (ca. 0.06—
0.13); propodeal spines long (ca. 0.20—
0.30); dorsum of head, except for the an-
tennal scrobes, with foveolate-reticulate
sculpture; postpetiole unsculptured; PPI
133-150; SL 0.70-0.82 and SI 81-93.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Worker. Tl 2.74—3.26, HL 0.78=0.92:
HW 0.80—0.96, CI 100-108, SL 0.70—0.82,
SI Si=93)) Ei O01 PW 0l50=0162%
AL 0.68—0. 20, BRW.0:23=02273RREOAG=
0.20, PPI 133-150 (@ = 74).
Mandibles usually with a few longitudi-
nal rugae. Masticatory margin of aancable
with three teeth: an apical + a preapical
+ a long diastema + a truncated basal
tooth. Basal margin of mandible lacking a
distinctly curved lobe or tooth. Clypeus
with a strong median carina. Anterior clyp-
eal margin with a median denticle and
usually two to three others on each side;
two or three lateral denticles are often
fused into one prominence. Ventral center
of clypeus with a low, broad, toothlike
prominence. Palp formula 2,3. Frontal ca-
rinae well developed, beyond the level of
the posterior margins of eyes. Scrobal ar-
eas shallow, present below the frontal ca-
rinae. Frontal lobes almost completely ab-
sent so that the antennal articulations are
entirely exposed. Antennal scapes, when
lying in the antennal scrobes, close to or
just reaching the occipital margin of head.
Eyes containing five to six Grae in
the longest row. Profile shape of alitrunk
and pedicel segments as in Figure 141.
Pronotum armed with a pair of short
spines (ca. 0.06—0.13). Propodeal spines
long, ca. 0.20 to 0.30, usually straight but
ROGUES slightly upcurved along their
length. Metapleural lobe small- triangular,
usually with an acute apex. Petiole hede in
profile higher than long, with a long an-
terior peduncle, its Lat ere dor sl angle
forming an apex and its dorsum sloping
downward posteriorly. Postpetiole in pro-
file much higher than long, with a rounded
dorsum; in der sal view, postpetiole trans-
verse-rectangled, much broader than long,
with the two sides subparallel. Dorsum of
head, except for the antennal scrobes
where there are only a few transverse ru-
gae, with well-developed foveolate-reticu-
late sculpture. Dorsum of alitrunk usually
with foveolate-reticulate sculpture and a
few coarse longitudinal rugae. Sides of
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
D»
&
447
Figures 140-143. Pristomyrmex foveolatus Taylor. 140A: Worker head, full-face view; 140B: Showing a toothlike prominence
on the center of ventral clypeus; 141: Worker, lateral view; 142A: Queen head, full-face view; 142B: Showing a toothlike prom-
inence on the center of ventral clypeus; 143: Queen, lateral view.
pronotum with a few foveolate punctures;
sides of the rest of alitrunk with some ir-
regularly superficial rugae. Petiole node
and postpetiole smooth and shining. Gas-
ter unsculptured. Dorsal surfaces of head
and alitrunk with numerous erect or sub-
erect long hairs. Dorsal surfaces of petiole
node and postpetiole with a pair of long
hairs, respectively, as shown in Figure 141.
First gastral tergite lacking erect or sub-
erect hairs. A few pairs of forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect to suberect short hairs. Color
reddish-brown.
Queen. TL 3.20, HL 0.83, HW 0.86, CI
104, SL 0.74, SI 86, EL 0.16, PW 0.66,
AL 0.87, PPW 0.25, PPL 0.16, PPI 139 (n
= Il),
General shape as in Figures 142-143,
with normal caste differences from the
conspecific worker; pronotum unarmed;
448
other characters similar to worker; propo-
deal spine length 0.25.
Male. Unknown.
Comments. Pristomyrmex foveolatus is
extremely similar to P. thoracicus, also
from Australia, in many characters in the
workers and queens, such as (1) the den-
tition of the masticatory margin of man-
dible, (2) palp formula, (3) structure and
shape of the clypeus, (4) length and shape
of the pronotal and propodeal spines, and
(5) sculpture of the dorsal surfaces of the
head and the alitrunk. The differences be-
tween the workers of these two species are
slight, as follows:
P. foveolatus
Antennal scapes shorter (SL 0.70—-0.82,
SI 81-93)
Postpetiole in dorsal view much broader
than long, PPI 133-150, with the two
sides subparallel, showing a_trans-
verse rectangle
P. thoracicus
Antennal scapes longer (SL 0.86—0.98,
SI 97-103)
Postpetiole in dorsal view slightly broad-
er than long, PPI 109-121, with the
two sides not subparallel, showing a
trapezoid
Without doubt, P. foveolatus and P. thor-
acicus are closely related to P. brevispino-
sus, from the Oriental region. The workers
of these three species all show the follow-
ing characters: (1) a pair of short pronotal
spines, (2) a toothlike prominence on the
ventral center of the clypeus, (3) a similar
structure and shape of petiole node in
both profile and dorsal views, (4) a similar
structure and shape of postpetiole in pro-
file, (5) well-developed foveolate-reticulate
sculpture on the dorsal head, and (6) dor-
sal alitrunk with a few coarse longitudinal
rugae. But P. foveolatus and P. thoracicus
can be separated from P. brevispinosus as
follows: The former two species have a
pair of long propodeal spines (ca. 0.19—
0.30), two segments of maxillary palpi, and
a truncated basal tooth on the masticatory
margin of mandible in the workers and
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
queens, while P. brevispinosus has a pair
of short propodeal spines or teeth (ca.
0.04—0.12), one segment of maxillary palp,
and usually two small basal teeth on the
masticatory margin of mandible.
The separation of both P. foveolatus and
P. thoracicus from the African P. cribrarius
(a member of the cribrarius group) is pro-
vided under P. cribrarius.
Material Examined (ANIC, MCZC).
Australia: N. Queensland, Clump Point,
rainforest floor, ex small wood fragment
(R. W. Taylor); Q., Clump Point, <20 m,
berlesate (Taylor and Feehan); N. Q., NW
of Daintree, Mt. Alexander, rainforest (P.
F. Darlington); Q., Alexandra Bay, 16.12°S,
145.26°E, rainforest, <50 m, berlesate
(Taylor and Feehan); Q. NE, Road, sum-
mit on Alexandra, Ra. Daintree, 16.15°S,
145.26°E, rainforest, 250 m,_ berlesate,
sieved litter (G. Monteith); Q., Kuranda,
Black Mt. Rd. 360 m, 17.47°S, 145.39°E,
rainforest, berlesate, sieved litter (G. Mon-
teith); Q., Kuranda, Black Mt. Rd. 430 m,
16.45 X 145.35, rainforest, berlesate (Tay-
lor and Feehan); Q., 4 km W of Kuranda,
450 m, 16.49 * 145.36, rainforest, berle-
sate (Taylor and Feehan); Q., 1 km W of
Kuranda, closed forest litter, berlesate (J.
Doyen); N Q., Kuranda, RF, in log (B. B.
Lowery); Q., NE, C. Tribulation, 16.08°S,
145.28°E, 20 m, rainforest, berlesate,
sieved litter (G. Monteith); Q., NE, Cape
Tribulation, Noah Ck, 5 m (G. B. Mon-
teith); NE Q., 1.5 km W of Cape Tribu-
lation (Site 3), 16.05°S, 145.28°E, 150 m,
rainforest, berlesate, sieved litter (Mon-
teith, Yeates, and Thomson); NE Q., 2.0
km WNW of Cape Tribulation (Site 2),
16.05°S, 145.28°E, 50 m, rainforest, ber-
lesate, sieved litter (Monteith, Yeates, and
Thomson); Q., near Cape Tribulation,
16.06°S, 145.28°E, 50 m, rainforest, ber-
lesate (Taylor and Feehan); Q., Thorton
Range, 16.14°S, 145.26°E, 100 to 150 m,
rainforest, berlesate (Taylor and Feehan);
Q., 20 km N of Cairns, rainforest (B. B.
Lowery); NE Q., Lyons Lookout, Rex
Hwy, Mossman, 400 m, rainforest, berle-
sate, sieved litter (G. Monteith and D.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 449
T\\
RW ER
SX
CoN
eae
\)
Figures 144-145. Pristomyrmex hirsutus sp. n. 144A: Worker head, full-face view; 144B: Showing a toothlike prominence on
the ventral clypeus; 145: Worker, lateral view.
Cook); NE Q., Hinchinbrook Is., Gayun-
dah Creek, 18.22°S, 146.13°E, 10 to 80 m,
rainforest, berlesate, sieved litter (Mon-
teith, Davies, Thompson, and Gallon); Q.,
NE, Mossman Gorge, 16.25°S, 145.20°E,
rainforest, berlesate, sieved litter (G. Mon-
teith); NE Q., Bakers Blue Mt., 17 km W
Mt. Molloy, 1,000 m, rainforest, berlesate,
sieved litter (G. Monteith and D. Cook):
Q., Mt. Cook, Nat. Pk., 15.29°S, 145.16°E,
rainforest, berlesate (A. Calder and J. Fee-
han); Q., Gap Creek, 15.50°S, 145.20°E, 5
km ESE of Mt. Finnigan, rainforest, ber-
lesate (A. Calder and J. Feehan); Q., N
Pingin Hill (J. Holt); Q., Mt. Windsor Ta-
bleland, ca. 850 m, 16.18°S, 145.05°E,
rainforest, berlesate (R. W. Taylor); N Q.,
28 km NNW Mt. Carbine, Windsor Table-
land, 900 m, rainforest, berlesate, sieved
litter (Monteith, Yeates, and Cook); Q.,
ME, Cannon Vale, 20.16°S, 148.43°E, 10
m, dry rainforest, berlesate, stick brushing
(G. Monteith); Q., ME, Mt. Dryander,
20.15°S, 148.33°E, 500 to 650 m, rainfo-
rest, berlesate, stick brushing (G. Mon-
teith); Q., ME, Brandy Ck Rd, Conway SF,
20.20°S, 148.42°E, 60 m, rainforest, ber-
lesate, sieved litter and stick brushing (G.
Monteith); Q., Finch Hatton Gorge,
21.04°S, 148.38°E, 470 m, mesophyll no-
tophyll vine forest, berlesate (A. Gillison);
OF Hinch Hatton) Gorges 21305 ¢ss
148.38°E, 200 m, rainforest, berlesate (R.
W. Taylor and T. A. Weir); N Q., 15.50°S,
145.20°E, 12-mi scrub Gap Creek, com-
plex mesophyll vine for. (Davies and Ra-
ven); N Q., 20 km N of Cairns, lowland
RF, creek between rocks (B. B. Lowery).
Ecological Information. This species oc-
curs only in rainforest in North Queens-
land, Australia, and has been collected in
litter berlesates; its nests are constructed
in rotting logs (Taylor, 1965).
Pristomyrmex hirsutus sp. n.
Figures 144-145
Diagnosis (Worker). Masticatory margin
of mandible with a long diastema; dorsal
head with a well-developed rugoreticulum;
petiole node in profile high, with a single
evenly blunt-rounded apex; first gastral
tergite with numerous erect or suberect
short hairs.
Holotype Worker (BMHH). TL 5.78,
Ee SS EW eles OleOle Seale COm Sil
A SEE OL2 SPW 090s Alea too RENV
0.38, PPL 0.42, PPI 90.
Mandibles smooth and shining. Masti-
catory margin of mandible with four teeth
arranged as two adjacent strong apical
teeth + a long diastema + two basal den-
ticles of similar size. Basal margin of man-
dible lacking a toothlike prominence.
Clypeus somewhat uneven, with a few
weak short rugae. Anterior clypeal margin
with a median denticle and two others on
each side (but one of them appears to be
fused by two small denticles). Frontal ca-
450
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 146-147. Pristomyrmex longispinus sp. n. 146:
rinae extending to the level of the poste-
rior margins of the eyes. Antennal scrobes
shallow, approximately ending at the level
of the posterior margins of eyes. Frontal
lobes weak so that the antennal articula-
tions are almost entirely exposed. Antennal
scapes long, when laid on the dorsal head,
surpassing Ache occipital margin by about
one-third of their length. eraaile shape of
alitrunk and pedicel segments as in Figure
145. Pronotum armed with a pair of spines
(ca. 0.19). Propodeum with a pair of some-
what elongate-triangular teeth (ca. 0. 09).
Metapleural lobes uben iangular. Petiole
node in profile high (ca. 0. 76), with a sin-
gle evenly blunt- rounded apex. Postpetiole
in profile rounded dorsally, in dorsal view
longer than broad and broadening from
front to back. Dorsum of head, except for
the scrobal areas, with well-developed
coarse rugoreticulum. Similar sculpture
present on the dorsum of the alitrunk and
the two sides of the pronotum. Sides of the
rest of the alitrunk with irregular coarse
rugae. Petiole, postpetiole, and gaster
SOoEl and shining. All dorsal surfaces of
body, including hea alitrunk, petiole
node, postpetiole, and gaster, with numer-
ous erect or suberect hairs, as shown in
Figure 145. A few pairs of forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
erect or suberect hairs. Color reddish-
brown.
Worker head, full-face view; 147: Worker, lateral view.
Queen and Male. Unknown.
Comments. Though only a single speci-
men is available so fir it undoubtedly rep-
resents a good species because it has nu-
merous erect or suberect hairs on the first
gastral tergite, which is possessed only by
fis new species in the Oriental fauna of
the quadridens group. In addition, this
species has an uneven clypeus and a dis-
tinct petiole node (bearing a single evenly
blunt-rounded apex) that is not seen in the
other Oriental species of the group.
Holotype Worker. Philippines: Misamis
or Mt. Balatukan, 15 km SW of Gingoog,
1,000 to 2,000 m, 1—5.v.1960 (H. Torrev-
illas).
Note: The holotype of this species is an
old specimen, and many hairs have obvi-
ously been removed from its first gastral
tergite. Therefore, the figure can aot ac-
curately show this character.
Ecological Information. Unknown.
Pristomyrmex longispinus sp. n.
Figures 146-147, 263, 272
Diagnosis (Worker). Frontal carinae
short, usually not extending to the level of
the posterior margins oe eyes; pronotal
spines exceptionally long (0. 42-0, 50); pro-
podeal spines moderately long (0. 18—0.26);
anterior face of petiole node, in profile, in-
distinguishable from the upper surface of
its anterior peduncle.
Holotype Worker (MCZC). TL5.30, HL
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
IRA SIV 23 Cll06, Siw 345 Si 1205s
EL 0.24, PW 0.80, AL 1.26. Paratypes, 73
workers and three males (BMNH, LACM,
MCZC, MHNG, USNM).
Worker. TL 4.52-5.62, HL 1.09—1.26,
EVV 15135) Cl 103=109) SE 126=1-40;
SI 103-113, EL 0.21—0.26, PW 0.76—0.86,
AL 1.14—1.32, PPW 0.31—0.35, PPL 0.36—
0.40, PPI 84—89 (n = 20).
Mandibles generally unsculptured,
smooth, and shining. Masticatory margin
of mandible with four teeth: the strongest
apical + the second strongest preapical +
a long diastema + two Saal teeth that are
roughly the same size. Basal margin of
mandible lacking a toothlike prominence.
Clypeus depressed, unsculptured, and
shining; its anterior margin with a median
denticle and usually two others on each
side, but sometimes two lateral denticles
are fused into one prominence. Ventral
center of clypeus with a short transverse
ruga or a broad-based weak prominence.
Palp formula 1,3. Frontal carinae short,
usually not extending to the level of the
posterior margins of the eyes. Antennal
scrobes absent. Frontal lobes very weak so
that the antennal articulations are almost
entirely exposed. Antennal scapes long,
when lying on the dorsal head, surpassing
the occipital margin by about one-fifth to
one-fourth of their length. Head in full-
face view subglobal. Profile shape of ali-
trunk and pedicel segments as in Figure
147. Pronotum armed with a pair of ex-
ceptionally long spines that are about 0.42
to 0.50 and longer than the distance be-
tween their bases. Propodeum with a pair
of moderately long spines that are 0.18 to
0.26 and over two times the distance be-
tween their bases. Both pronotal and pro-
podeal spines acute and directed upward.
Metapleural lobes subtriangular. Dorsum
of alitrunk sometimes slightly concave. An-
terior face of petiole node, in profile, in-
distinguishable from the dorsal surface of
its anterior peduncle (i.e., the anterior face
of petiole node and the dorsal surface of
peduncle forming a long declivity from the
base of peduncle to the top of petiole
451
node). Postpetiole in profile rounded dor-
sally, in dorsal view longer than broad.
Dorsum of head generally smooth and
shining, but a few weak short rugae pres-
ent on the frontal area and sometimes a
few foveolate punctures on the genae. Al-
itrunk smooth and polished. Petiole, post-
petiole, and gaster unsculptured, smooth,
and shining. Dorsal surfaces of head and
alitrunk with some erect or suberect long
hairs. Usually two to three pairs of snag
hairs present on the dorsal surfaces of pet-
iole node and_ postpetiole, respectively.
First gastral tergite usually lacking erect or
suberect hairs, rarely with few hairs. A few
pairs of forward-projecting hairs present
near the anterior clypeal margin. Scapes
and tibiae with some erect or suberect
hairs. Color reddish-brown.
Queen. Unknown.
Male (Figs. 263, 272). Three male spec-
imens, together with 48 workers, collected
in Dumaguete, Philippines, by J. W. Chap-
man, Gonstiuite a series; and two of the
three males were originally mounted, re-
spectively, with a worker on the same pin:
TL 4.42-4.54, HL 0.67-0.73, HW 0.64—
0.66, CI 88-96, SL 0.20-0.22, SI 31-34,
HWE 0.94—0.98, EL 0.43—-0.46, PW 0.8S—
0.93, AL 1.32-1.44, PPW 0.28-—0.30, PPL
0.242026. PP 5=1l7 Gi — 3):
Head, including the eyes, distinctly
broader than long. “Clypeus convex, with-
out a median longitudinal carina. Palp for-
mula 1,3. F rental carinae sometimes ab-
sent but sometimes present, just reaching
the level of the posterior margins of an-
tennal insertions. Maximum length of the
median ocellus 0.12 to 0.12. On the me-
sonotum, notauli pronounced, forming a Y
shape; parapsidal furrows absent. Scuto-
scutellar sulcus with 12 to 13 narrow ridg-
es. Middle and hind tibiae without any
spurs. Propodeum slightly tuberculate,
lacking spines and teeth. Metapleural
lobes prominent and subtriangular. Petiole
node in profile low, with a subtriangular
apex and a rather long anterior peduncle;
anterior face of the Rode together with
the dorsal surface of the peduncle, form-
452
ing a long declivity that reaches the top of
the node. Postpetiole in profile low, round-
ed dorsally, and in dorsal view broader
than long. Dorsum of head smooth and
shining, but frontal area usually with a me-
dian longitudinal carina. Alitrunk smooth
and shining, except for those marked su-
tures. Petiole, postpetiole, and gaster un-
sculptured, smooth, and shining. All dorsal
surfaces with abundant erect or suberect
hairs. Scapes and tibiae with numerous
erect or suberect short hairs. Color red-
dish-brown; hairs reddish-brown; wings
somewhat infuscated.
Comments and Discussion. Pristomyr-
mex longispinus is closely related to P. cur-
vulus. The former is very similar in the
workers to the latter in the shape of pro-
notal and propodeal spines as well as in
the size, sculpture, hair, and color of body.
The differences between the two species
are provided under P. curvulus.
Pristomyrmex longispinus may have
evolved from a P. flatus—like ancestor. Ex-
cept for the well-developed pronotal
spines, its larger size, and shorter frontal
carinae, the workers of P. longispinus are
similar to those of P. flatus.
It is possible that a P. collinus—like an-
cestor may have split into the four species
P. collinus, P. flatus, P. curvulus, and P. lon-
gispinus because morphological characters
show that (1) P. collinus and P. flatus are
a pair of sibling species, (2) P. curvulus and
P. longispinus are another pair of sibling
species, and (3) PB. curvulus seems to be
derived from a P. collinus—like ancestor
(see the discussion under P. curvulus).
This hypothesis also obtains support from
biogeographic data. The four species are
all endemic to the Philippines. Pristomyr-
mex curvulus and P. longispinus are found
only in Dumaguete and P. flatus only in
Luzon. But P. collinus has a larger range;
it occurs sympatrically with P. curvulus and
P. longispinus in Dumaguete and with P.
flatus in Luzon.
Holotype Worker. Philippines: Duma-
guete (J. W. Chapman).
Paratypes. Philippines: 48 workers and
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
three males with same data as holotype;
four workers, Dumaguete, Horns of Ne-
gros (J. W. Chapman); two workers, Du-
maguete, 1942 (J. W. Chapman); one
worker, Dumaguete, 7.xi.43 (J. W. Chap-
man); two workers, Dumaguete, v.1947 (J.
W. Chapman); one worker, Dumaguete,
13.v.1947 (J. W. Chapman); one worker,
Dumaguete, Silliman University, 1948
(Domingo Empeso); 13 workers, Duma-
guete, 1949 (J. W. Chapman); one worker,
Dumaguete, 18.vi.49. (J. W. Chapman).
Additional Material Examined. More
than two dozen specimens from Camp,
Dumaguete, Philippines, are treated as
non-type material because of in a poor sit-
uation.
Ecological Information. Unknown.
Pristomyrmex modestus sp. n.
Figures 148-149
Diagnosis (Worker). Pronotum with a
pair of moderately long, robust spines;
dorsal surfaces of head and _ alitrunk, as
well as the two sides of pronotum, with a
coarse rugoreticulum; petiole node in pro-
file somewhat transrectangular, slightly
longer than high, with seven to eight fo-
veolate punctures.
Holotype Worker (BMNH). TL 4.00,
HL 0.99, HW 0.92, CI 93, SL 0.95, SI 103,
EL O19) PW 0.64-AL 104) RPRW.0)35%
PRIEO:355 REIOO!
Mandibles with a few longitudinal ru-
gae. Masticatory margin of mandible with
two adjacent strong apical teeth + a long
diastema + one broad, somewhat concave
basal tooth. Basal margin of mandible lack-
ing a distinctly curved lobe or tooth. Clyp-
eus with a strong median carina. Anterior
clypeal margin with a median denticle and
three others on each side. Ventral surface
of clypeus with a short transverse ruga.
Frontal carinae strong, extending to the
level of the posterior margins of the eyes. |
Antennal scrobes indistinct. Frontal lobes |
weak so that the antennal articulations are
almost entirely exposed. Antennal scapes,
laid on the dorsal head, slightly surpassing _
the occipital margin of head. Eyes rather —
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
453
Figures 148-149. Pristomyrmex modestus sp. n. 148A: Worker head, full-face view; 148B: Showing a short transverse ruga
on the ventral clypeus; 149: Worker, lateral view.
large. Profile shape of alitrunk and pedicel
segments as in Figure 149. Pronotal spines
robust, ca. 0.14, shorter than the distance
between their bases. Propodeal spines
acute, slender, ca. 0.11. Metapleural lobes
developed, prominent, ca. 0.14, each with
a rounded apex. In profile view, petiole
node slightly longer than high, somewhat
transrectangular, with the anterodorsal an-
gle on approximately the same level as or
weakly higher than the posterodorsal; in
dorsal view, petiole node longer than
broad. Postpetiole in profile rounded dor-
sally; in dorsal view, approximately quad-
rate and about as long as broad. Dorsal
surfaces of head and alitrunk, as well as
the two sides of pronotum, entirely sculp-
tured with coarse rugoreticulum. Dorsum
and the sides of petiole node with seven
to eight large foveolate punctures. Post-
petiole with a few shallow foveolate punc-
tures. Gaster unsculptured, smooth, and
shining. Dorsal surfaces of head and ali-
trunk with numerous erect or suberect
hairs. Petiole node and postpetiole each
with a few pairs of hairs, as shown in Fig-
ure 149. First gastral tergite lacking erect
or suberect hairs. A few pairs of forward-
projecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
sparse erect or suberect hairs. Color red-
dish-brown.
Queen and Male. Unknown.
Comments. This species must have
evolved from the ancestor of P. costatus. It
differs from P. costatus in the workers as
follows:
P. modestus
Petiole node in profile longer than high,
somewhat rectangular, with the anter-
odorsal angle on approximately the
same level as the posterodorsal
Dorsum and sides of petiole node with
seven to eight foveolate punctures
P. costatus
Petiole node in profile higher than long;
its anterodorsal angle is distinctly
higher than the posterdorsal
Dorsum and sides of petiole node with-
out foveolate punctures
Holotype Worker. Malaysia: Sarawak,
4th Division, Gn. Mulu N.P., Kerangas
for., leaf litter, 19.iii.1978 (H. Vallack).
Ecological Information. The holotype is
collected from a forest.
Pristomyrmex nitidissimus Donisthorpe
Figures 150-151
Pristomyrmex nitidissimus Donisthorpe, 1949: 411.
Holotype worker, New Guinea: Maffin Bay, ix.1944
(E. S. Ross) (CASC) [examined].
454
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 150-151.
ruga on the ventral clypeus; 151: Worker, lateral view.
Diagnosis (Worker). Pronotum armed
with a pair of teeth; dorsal surfaces of head
and alitrunk with numerous scattered fo-
veolate punctures; ventral surface of clyp-
eus with a coarse transverse carina; larger
size (HL 1.10=1.16, HW 1.22-1.24 and
EE tOh24=022'5))
Worker. TL 4.58, 4.69: HL 1.10, 1.16:
FIV 2 2124 CialO al Sipe A eG:
SI 93, 94: EL. 0.24, 0.25: PW 0.75, 0.78;
IN IAD TO) Ge SD).
Mandibles with a few longitudinal ru-
gae. Dentition of the masticatory margin
of mandible: the strongest apical + the
second strongest preapical + a short dia-
stema + a broad basal tooth showing two
minute points (which is formed by the fu-
sion of two basal denticles). Basal margin
of mandible with a central, broadly curved
lobe. Clypeus with a median longitudinal
carina. Anterior clypeal margin with a me-
dian denticle and two others on each side.
Ventral surface of clypeus with a trans-
verse ridge. Frontal carinae extending to
the level of the posterior margins of eyes.
Antennal scrobe indistinct, but a smooth
area present below the frontal carina.
Frontal lobes very weak so that the anten-
nal articulations are almost entirely ex-
posed. Antennal scapes, laid on the dorsal
head, slightly surpassing the occipital mar-
gin of head. Eyes large. Pronotum armed
with a pair of teeth. Propodeum with a
Pristomyrmex nitidissimus Donisthorpe. 150A: Worker head, full-face view; 150B: Showing a short transverse
pair of subtriangular short spines that are
slightly longer than the pronotal teeth.
Metapleural lobes each with a subtrian-
gular apex. Petiole node in profile as in
Figure 151, with a fairly long anterior pe-
duncle, in dorsal view longer than broad.
Postpetiole in profile rounded dorsally, in
dorsal view slightly longer than broad.
Dorsum of head with numerous rather
large, scattered foveolate punctures; space
between foveolae usually smooth. Similar
foveolate punctures present on the dorsal
surface of alitrunk, but promesonotum
with a smooth, unsculptured median strip.
Petiole, postpetiole, and gaster smooth
and shining. Dorsal surfaces of head, ali-
trunk, petiole node, and postpetiole with
numerous erect or suberect hairs. First
gastral tergite lacking erect or suberect
hairs. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with some erect
to suberect short hairs. Color blackish-
brown.
Queen and Male. Unknown.
Comments. At first glance, P. nitidissi-
mus appears to resemble P. quadridens
very much, but after being compared in
detail, the workers of the two species are
separable as follows:
P. nitidissimus
Ventral surface of clypeus with a coarse
transverse ruga, lacking a toothlike |
prominence
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
<i
Ly
VY B
455
Figures 152-153. Pristomyrmex occultus sp. n. 152A: Worker head, full-face view; 152B: Showing a tooth on the center of
ventral clypeus; 153: Worker, lateral view.
Larger species, with HW 1.22—1.24, HL
OS Ga Ele Oh 420825
Basal margin of mandible with a central,
broadly curved lobe
Four to five pairs of short hairs present
on the dorsums of both petiole node
and postpetiole
P. quadridens
Ventral center of clypeus with a tooth-
like prominence
Smaller species, with HW _ 0.82-1.02,
UE OlS2 =I" 023 Ee Oni4z=020
Basal margin of mandible almost
straight, without a distinctly convex
lobe
Usually one to two pairs of hairs present
on the dorsums of both petiole node
and postpetiole
The workers of P. nitidissimus are ob-
viously different from those of the African
P. africanus because the latter are smaller
(HW < 1.00, HW < 1.00, EL < 0.14) and
lack hairs on the dorsal surfaces of the pet-
iole node and the postpetiole. The workers
of P. africanus also possess two toothlike
prominences on the ventral surface of the
clypeus and have a concave dorsum of the
alitrunk and an almost straight basal mar-
gin of the mandible.
Material Examined (CASC). One work-
er, with same data as holotype: New Guin-
ea: Maffin Bay, ix.44 (E. S. Ross).
Ecological Information. Unknown.
Pristomyrmex occultus sp. n.
Figures 152-153
Diagnosis (Worker). Masticatory margin
of mandible with five teeth; pronotum
with a pair of triangular short spines; dor-
sal surfaces of head and alitrunk, as well
as the two sides of pronotum, with coarse
rugoreticulum.
Holotype worker (BMNH). TL 3.10, HL
0.84, HW 0.81, CI 96, SL 0.72, SI 89, EL
0.10, PW 0.56, AL 0.80. Paratypes, three
workers (MHNG, MCZC).
Worker. TL 3.04—3.23, HL 0.84—0.88,
HW 0.80-0.84, CI 93-98, SL 0.72-0.75,
S693) EE O0=0nl35 RW 0:52=0'56:
AL 0.76—0.80, PPW 0.22-0.24, PPL 0.20—
0.22, PPI 109-120 (n = 5).
Mandibles generally smooth and _ shin-
ing, with a few basal longitudinal rugae.
Masticatory margin of mandible with five
teeth arranged as the strongest apical +
the second strongest preapical + a diaste-
ma + three small denticles of similar size
of which the middle one is sometimes
weak (i.e., smaller than the two others) or
worn down, but the length of the masti-
catory margin covered by the three den-
456
ticles is longer than the diastema. Basal
margin of mandible lacking a toothlike
prominence. Clypeus depressed, smooth,
and shining. Anterior clypeal margin with
a median denticle and two to three others
on each side. Ventral center of clypeus
with a strongly prominent tooth. Frontal
carinae strong, extending to the level of
the posterior margins of eyes. Antennal
scrobes shallow. F rental lobes weak; thus,
the antennal articulations are almost en-
tirely exposed. Antennal scapes, laid on the
dorsal head, just reaching to the occipital
margin of head. Eyes moderately sized,
containing five to six ommatidia in the lon-
gest row. Occipital margin in full-face view
feebly concave. Pronotum armed with a
pair of triangular short spines. Propodeal
spines acute, about two times the length
of the pronotal teeth. Metapleural lobes
prominent and rounded. Dorsum of ali-
trunk in dorsal view somewhat depressed.
Petiole node high in profile, with a fairly
long anterior peduncle; its anterodorsal
angle higher than the posterodorsal. Sub-
petiole with a narrow, long, semitranslu-
cent lamella. Postpetiole in “profile round-
ed dorsally, in dorsal view slightly broader
than long. Dorsal surfaces of head and al-
itrunk, as well as the two sides of prono-
tum, with coarse, strongly sculptured ru-
goreticulum, but the scrobal areas lacking
this sculpture. Petiole and postpetiole
smooth and shining, except for a lateral
longitudinal carina on each side that sep-
arates the tergite from the sternite. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with numerous erect or suberect
hairs. Two pairs of hairs present on the
dorsum of petiole node and usually two to
three pairs on the dorsum of postpetiole.
First gastral tergite lacking erect or sub-
erect hairs. A few of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with some erect
to suberect short hairs. Color reddish-
brown.
Queen and Male. Unknown.
Comments. Pristomyrmex occultus is a
sibling species of P. quindentatus. The two
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
species are all from the Oriental region.
The separation of the two species is dis-
cussed under P. quindentatus.
Pristomyrmex occultus is also very sim-
ilar in appearance to P. brevispinosus. But
in the workers of P. occultus, the mastica-
tory margin of the mandible possesses five
teeth, the length of the masticatory margin
covered by three small denticles is slightly
longer than that of diastema, and the pro-
podeum is armed with a pair of fairly long
spines. In PB. brevispinosus, the masticatory
margin of the mandible has four teeth, the
length of the masticatory margin covered
by two basal denticles is not longer than
that of diastema, and the propodeum is
armed with a pair of triangular teeth.
Holotype Worker. E. Malaysia: Sarawak,
confl. Sun Oyan and Mujong riv., E Kapit
50 m, 18.v.1994 (L6bl and Burchhardt).
Paratypes. Three workers, Malaysia: Sa-
bah, Poring Hot Springs, 600 m, 9.v.1987
(Burckhardt and Lo6bl).
Additional Material Examined. Two
workers (BMNH, MCZC), collected in
Malaysia (Sabah, Poring Hot Springs, 500
m), by Burckhardt and L6bl on 7.v.1987,
show only four distinct teeth on the mas-
ticatory margin of the mandible. But the
length of the masticatory margin covered
by two small basal denticles is slightly lon-
ger than that of diastema.
Ecological Information. Unknown.
Pristomyrmex orbiceps (Santschi)
Figures 154-157, 264, 273
Xiphomyrmex orbiceps Santschi, 1914: 367. Syntype
workers, Cameroon: Victoria (F. Silvestri); and
Ghana: Aburi (F. Silvestri) (NHMB) (Bolton,
1981).
Pristomyrmex orbiceps (Santschi) Santschi, 1916: 51.
Hylidris laevigatus Weber, 1952: 20. Holotype work-
er, Zaire: 13 mi S of Asa, lat. 4°40’N, long. 25°40'E
(N. A. Weber) (AMNBH). [Synonymy by Bolton,
1981].
Note: I have not seen the type material,
but I did view some specimens of this spe-
cies, which has been compared with the
type by B. Bolton.
Diagnosis (Worker). Masticatory margin —
of manchble with a diastema, after the —
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
457
Figures 154-157. Pristomyrmex orbiceps (Santschi). 154A: Worker head, full-face view; 154B: Showing a transverse ruga on
the ventral clypeus; 155: Worker, lateral view; 156A: Queen head, full-face view; 156B: Showing a transverse ruga on the ventral
clypeus; 157: Queen, lateral view.
preapical tooth; pronotum lacking teeth or
spines; dorsal surfaces of head between
the frontal carinae and alitrunk unsculp-
tured, smooth, and shining; petiole and
postpetiole lacking erect or suberect hairs.
Worker. TL 2.66-3.40, HL 0.78-0.92,
HW 0.80-0.98, CI 100—110, SL 0.68—0.80,
SI 79-85, EL 0.14—0.21, PW 0.51-0.60,
AL 0.68—0.80, PPW 0.24—0.30, PPL 0.1S5—
0.22, PPI 125-144 (n = 35).
Mandibles smooth, at most with two
longitudinal basal rugae. Masticatory mar-
gin of mandible with an apical tooth + a
preapical tooth + a diastema + a broad,
truncated (or sometimes somewhat mid-
concave) basal tooth. Basal margin of man-
dible lacking a toothlike prominence or
curved lobe. Clypeus possessing or lacking
a longitudinal median carina. Ventral sur-
face of clypeus with a long transverse
ridge. Anterior clypeal margin with a me-
dian denticle and two to three others on
each side, but some of the denticles are
usually vestigial or very weak; sometimes,
the margin without any distinct denticles.
Palp formula 1,3. Frontal carinae extend-
ing to the level of the posterior margins of
eyes. Antennal scrobes absent. Frontal
lobes absent. Antennal scapes, when lying
on the dorsal head, close to the occipital
margin of head. Eyes containing six to nine
ommatidia in the longest row. Pronotum
458
tuberculate, lacking teeth or spines. Pro-
podeum armed with a pair of triangular
teeth or short spines. Metapleural lobes
rounded. In profile view, petiole node
massive, higher than long, with a robust
anterior peduncle; its anterodorsal and
posterodorsal angles rounded, but the for-
mer is higher than the latter (Fig. 155). In
dorsal view, petiole node slightly broader
than long. Postpetiole in profile higher
than long, rounded dorsally, in dorsal view
broadening from front to back and dis-
tinctly broader than long. Dorsum of head
between the frontal carinae smooth and
shining. A few foveolate punctures present
on thee sides of the dorsal head and some-
times bordering the frontal carinae. Ali-
trunk, petiole, postpetiole, and gaster un-
sculptured, smooth, and shining. Dorsum
of head beyond the level of antennal in-
sertions with about four pairs of hairs:
three pairs on the frontal carinae and one
pair on the occipital corners. Dorsal sur-
face of alitrunk with only a pair of hairs
present on the mesonotum. Petiole, post-
petiole, and first gastral tergite lacking
erect or suberect hairs. Several pairs of
forward-projecting hairs present near the
anterior clypeal margin. Scapes and tibiae
with appressed hairs. Color blackish-
brown.
Queen. IL, SOL B58) IeUk, Osea =O.80),
HW 0.91—0.99, CI 106-110, SL 0.74—0.79,
SI80=85) JE 1 0!23=0265 PW. 017 020577.
AL 0.90—1.00, PPW 0.29-—0.32, PPL 0.2]—
0.24, PPI 129-145 (n = 8).
General shape as in Figures 156-157,
with normal caste differences from con-
specific worker; eyes larger; mesonotum
with more eure than in the conspecific
worker. Other characters similar to worker.
Male. (Figs. 264, 273). Two male speci-
mens, one collected in Ghana (Tafo, twig
in leaf litter), by B. Bolton, and the second
male in Ivory Coast (Orston Expt. Sta., 17
km. W. of Abidjan), by W. L. Brown, were
originally mounted, respectively, together
ih one and two workers on hes same
pins: TL 2.58, 2.76; HL 0.48, 0.50; HW
0.47, 0.48; CI 96, 100, SL 0.18, 0.18; SI
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
38, 38; HWE 0.63, 0.66; EL 0.24, 0.26:
PW 0.57, 0.58; AL 0.87, 0.92; PPW 0.20,
Oils IIB Oe, O.lles IPP WL, WIS Ge =
2).
Head, including the eyes, distinctly
broader than long. Clypeus narrow, convex
in middle; antenion clypeal margin trans-
verse. Maximum diameter of the median
ocellus 0.08 to 0.09. On the mesoscutum,
notauli distinct, forming a Y shape; par-
apsidal furrows absent. Scuto-scutellar sul-
cus with 8 to 11 short ridges. Propodeum
slightly tuberculate, lacking teeth or
spines. Metapleural lobes subtriangular.
Petiole node low in profile, with a fairly
long anterior peduncle; its anterior face,
together with the dorsal surface of the pe-
duncle, forming a declivity. Postpetiole in
profile low, rounded dorsally, and in dorsal
view slightly broader than long. Dorsum of
head generally smooth and shining, but
with about 8 to 10 short rugae on the pos-
terior clypeal margin. A few short rugae
present on the frontal area. A weak,
curved ruga present above the antennal in-
sertion. Nira generally smooth and
shining, except for those marked sutures.
Petiole, postpetiole, and gaster smooth
and shining. All dorsal siuhiges with abun-
dant erect or suberect hairs. Scapes and
tibiae with numerous suberect short hairs.
Body reddish-brown; wings white or light-
yellow.
Comments and Discussion. Pristomyr-
mex orbiceps occurs in Africa. It is some-
what similar, in appearance of the workers,
to P. fossulatus, a member of the punctatus
group, also from Africa. The workers of
the two species all lack pronotal teeth or
spines. But they can be separated as fol-
lows: (1) P. orbiceps possesses a single seg-
ment of maxillary palp, in contrast with
four segments in P. fossulatus; (2) P. fos-
sulatus has a wedge-shaped petiole node
that is not seen in P. orbiceps; and (3) the
dorsal surfaces of the head and the ali-
trunk are smooth in P. orbiceps but sculp-
tured with scattered foveolate punctures in
P. fossulatus.
Pristomyrmex orbiceps belongs to the |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 459
rainforest, Berlesate (R. W. Taylor); Tafo,
quadridens group. The absence of the pro-
notal teeth or spines in the workers should
not impede this species from being as-
signed to the group. P. africanus, a distinct
member of the quadridens group, some-
times also possesses only a pair of tuber-
cles on the pronotum, like P. orbiceps.
Pristomyrmex orbiceps is easily recog-
nized in the quadridens group. It differs
from the other species of the group, except
for P. eduardi and sometimes P. africanus,
in the workers in lacking a pair of pronotal
teeth or spines. Pristomyrmex orbiceps dif-
fers from P. eduardi and P. africanus in the
workers as follows: The masticatory mar-
gin of the mandible possesses five teeth in
P. eduardi but three teeth in P. orbiceps;
the eyes contain three to four ommatidia
in the longest row in P. eduardi and four
to five ommatidia in P. africanus but six to
nine ommatidia in P. orbiceps; the dorsal
surfaces of the head and the alitrunk are
smooth in P. orbiceps but sculptured with
scattered foveolate punctures in P. african-
us and with rugoreticulum in P. eduardi;
the ventral surface of the clypeus bears a
central tooth in P. eduardi, two prominent
teeth in P. africanus, but a long transverse
ridge in P. orbiceps; the dorsum of the al-
itrunk has only one pair of hairs in P. or-
biceps but numerous in both P. eduardi
and P. africanus.
Distribution. Ivory Coast, Ghana, Ni-
geria, Cameroon, Gabon, and Angola (Bol-
ton, 1981). The following records (MCZC,
ANIC) are added here: Angola: Dundo,
Carnisso, barks eRe sLwachime. 17.2228.
20.50°E, gallery forest, berlesate (native
collector); Dundo, R. Luachimo (Rte. Tur-
ismo), 7.025°S, 20.51°E, gallery forest, ber-
lesate ates collector). Ivory Coast: Divo,
rainforest, litter (L. Brader); Gagnoa, rain-
forest, litter (L. Brader). People’s Rep.
Congo: 25 k NW Boha, 30 k SE Lac Telle
(Gary Alpert). French Equatorial Africa:
Ubangi-Shari, Bas Mbomu (N. A. Weber).
Ghana: E.R. Mt. Atewa, rainforest, Ber-
lesate (R. W. Taylor); E.R. Scarp Forest,
near Bosuso, rainforest, Berlesate (R. W.
Taylor); E.R. Nkwanda For., near Enyiresi,
Eastern Reg., rainforest, Berlesate (R. W.
Taylor).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter berlesates (also see Weber, 1952:
2)
Pristomyrmex quadridens Emery
Figures 158-161, 274
Pristomyrmex quadridens Emery, 1897: 584. Lecto-
type worker, New Guinea: Friedrich-Wilhelmshaf-
en et Berlinhafen (L. Biré) (MCSN), here desig-
nated, [examined].
Pristomyrmex quadridens var. aruensis Karavaiev,
1933: 270. Holotype worker, Indonesia: Aru Is.,
Wammar (W. Karawajew) (UENC) [examined].
Syn. n.
Pristomyrmex orbiculatus Donisthorpe, 1948: 306.
Syntype workers, queens and males, New Guinea:
Maffin Bay, 20.vi.1944 (E. S. Ross) [syntype work-
ers (BMNH, LACM, USNM) examined]. Syn. n.
Diagnosis (Worker). Pronotum and pro-
podeum each with a pair of short spines;
dorsal surfaces of alitrunk and head be-
tween the frontal carinae with scattered
foveolate punctures; petiole node and
postpetiole each with at least a pair of
hairs; HW 0.82—1.02, HL 0.82-1.02.
Worker Tl 334-410) Ale 078221202)
HW 0.82-1.02, CI 96-102, SL 0.86—1.00,
SI 96-105, EL 0.14—0.20, PW 0.54—0.70,
AL 0.82-1.04, PPW 0.24—0.28, PPL 0.26—
0.30, PPI 86—L00 (n = 30).
Mandibles usually smooth and shining
but sometimes with a few fine longitudinal
rugae. Dentition of the masticatory margin
of mandible: the strongest apical tooth +
the second strongest preapical + a long
diastema + two Srna teeth that are rough
ly the same size; sometimes two small bas
al teeth are fused, forming a broad tooth
with two points. Basal margin of mandible
lacking a distinct toothlike prominence.
Clypeus depressed and smooth, with a me-
dian longitudinal carina, but sometimes
this raeclon carina weak or absent. Ante-
rior clypeal margin usually with a median
denticle and two to three others on each
side. Ventral surface of clypeus with a
toothlike prominence at the center. Palp
460
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 158-161.
Pristomyrmex quadridens Emery. 158A: Worker head, full-face view; 158B: Showing a tooth on the center
of ventral clypeus; 159: Worker, lateral view; 160A: Queen head, full-face view; 160B: Showing a tooth on the center of ventral
clypeus; 161: Queen, lateral view.
formula 1,3. Frontal carinae distinct, ex-
tending to the level of the posterior mar-
gins of eyes. Antennal scrobes shallow and
short. Frontal lobes weak; thus, the anten-
nal articulations are almost entirely ex-
posed. Antennal scapes, when lying on the
dorsal head, slightly surpassing the occip-
ital margin of head. Eyes usually contain-
ing seven to eight ommatidia in the longest
row. Occipital margin feebly concave. Pro-
file of alitrunk and pedicel segments as in
Figure 159. Pronotum armed with a pair
of acute short spines. Propodeum with a
pair of triangular teeth or short spines that
are about equal in length to the pronotal
armaments. Metapleural lobes rounded or
with a subtriangular apex. Petiole node in
profile nodiform; its anterodorsal angle
high, and the dorsum sloping downward
posteriorly. Postpetiole in profile rounded
dorsally, in dorsal view broadening from
front to back and usually longer than
broad but sometimes about as long as
broad. Dorsum of head between the fron-
tal carinae with scattered foveolate punc-
tures; intensity and number of punctures
very variable: The punctures are some-
times dense and large but sometimes few,
small, and shallow. Dorsum of alitrunk
with scattered foveolate punctures, varying |
from a few to many. Petiole, postpetiole,
and gaster unsculptured, smooth, and
shining. Dorsal surfaces of head and ali-
trunk with numerous erect or suberect |
hairs. Two pairs of hairs usually present on
the dorsum of petiole node and a pair usu-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
ally on the dorsum of postpetiole. First
gastral tergite without erect or suberect
hairs. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with abundant
erect to suberect short hairs. Color varying
from light yellow-brown to dark-brown; in
some specimens, a darker circular patch
present on the dorsum of head just above
the frontal area; sometimes gaster darker
than head and alitrunk.
Queen. TL 3.82-4.72, HL 0.90-1.10,
HW 0.94—1.11, CI 98-105, SL 0.88—1.02,
SI 90-99, EL 0.23—0.28, PW 0.74—-0.88,
AL 1.06—1.26, PPW 0.26—0.32, PPL 0.29—
O34. PPI 88=97 (= 7):
General shape as in Figures 160-161,
with normal caste differences from the
conspecific worker; pronotum unarmed.
Other characters similar to worker.
Male (Fig. 274). Two male specimens,
together with 12 workers and two queens,
constitute a series collected in Indonesia
(Seram, above Haruru, near Masohi, 50—
150 m) by W. L. Brown; each of the two
males was originally mounted with a work-
er on the same pin: TL 3.30, 3.52; HL
0.60, 0.62; HW 0.60, 0.62; CI 100, 100; SL
0.23, 0.24: SI 38, 39; HWE 0.80, 0.82; EL
0.32, 0.33; PW 0.79, 0.82; AL 1.20, 1.20:
BREW e022 5 0922] PPIi 025.) O27 Rel sie
83 G@ =
Head, including the eyes, distinctly
broader than long. Clypeus somewhat
transverse, convex, and unsculptured. Palp
formula 1,3. Frontal carinae absent. Max-
imum diameter of the median ocellus 0.11
to 0.12. On the mesoscutum, notauli pro-
nounced, forming a Y shape; parapsidal
furrows very weak. Scuto-scutellar sulcus
with 10 narrow longitudinal ridges. Pro-
podeum tuberculate, lacking teeth or
spines. Metapleural lobes somewhat
rounded. Petiole node in profile low and
rounded dorsally, with a fairly long ante-
rior peduncle. Postpetiole low, in dorsal
view distinctly longer than broad. Dorsum
of head smooth and shining. Alitrunk
smooth and shining, except for those
marked sutures. Petiole, postpetiole, and
461
gaster smooth and shining. All dorsal sur-
faces with abundant long hairs. Scapes and
tibiae with numerous erect or suberect
short hairs. Color blackish-brown; hairs
reddish-brown; wings somewhat infuscat-
ed.
Comments and Discussion. This species
occurs in New Guinea, some islands near
New Guinea, and Pohnpei Island. One of
its closely related species, P. brevispinosus,
is distributed in Indonesia, Malaysia, Thai-
land, the Philippines, Taiwan, and Japan;
in other words, P. brevispinosus occurs to
the northwest of P. quadridens (see Fig.
162). The differences between the workers
of the two species are as follows: The dor-
sal surfaces of the head and the alitrunk
possess scattered foveolate punctures in P.
quadridens but foveolate-reticulate sculp-
ture or rugoreticulum in P. brevispinosus.
Characters separating P. quadridens
from P. nitidissimus of New Guinea are
presented under P. nitidissimus. A discus-
sion of separating P. quadridens from the
Asian P. collinus and P. flatus is provided
under P. flatus. The separation of P. quad-
ridens from the African P. africanus is giv-
en under the latter name. The three Aus-
tralian species, P. wheeleri, P. erythropy-
gus, and P. quadridentatus, differ from P.
quadridens in the workers in having three
strong teeth on the anterior clypeal mar-
gin, three teeth on the masticatory margin
of the mandible, longer pronotal arma-
ments, palp formula of 2,2, and lacking fo-
veolate punctures on the dorsal surfaces of
the head and the alitrunk.
Material Examined (MCZC, BMNH,
ANIC, LAMN, USNM, NHMV, BMHE#).
Papua New Guinea: Gulf Prov., Ivimka
Camp, Lakekamu Basin, 7.7°S, 146.8°E,
120 m el, lowland wet forest, #96-205,
#96-232 (nest in rotting tree limb), #96-
234 (misc. strays), #96-300 (in rotten
branch on ground), #96-330 (foragers on
log) (R. R. Snelling); N. Dist., Sangara (P.
M. Room); N. Dist., 12 m N Popondetta
(B. B. Lowery); Brown R., rainforest, low-
land (B. B. Lowery); Bewani Rd. near Van-
imo ca. 7 to 10 km, 240 to 380 m, rain-
462
@ &. brevispinosus
A P. quadridens
Figure 162.
forest (W. L. Brown); ca. 12 km SE Vani-
mo, virgin hill, rainforest, 150 m (W. L.
Brown); Nadzab, dry evergreen forest (E.
O. Wilson); Huon Peninsula, Lower Busu
R., lowland rainforest (E. O. Wilson); near
Dobodura, Samboga R. 400 ft, rainforest
(B. B. Lowery); Bulolo 2600 ft, rainforest
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
gl
Distributions of Pristomyrmex brevispinosus and Pristomyrmex quadridens.
(B. B. Lowery); 12 mi N of Popondetta,
Bisicocoa Plan., 400 ft, rainforest (B. B.
Lowery); Lae, Busu R. area, rainforest (B.
B. Lowery); Lae, Markham R. Bridge, —
rainforest (B. B. Lowery); near Lae, <50
m (R. W. Yaylor); ca. 16 km: NW Laew
“Timber Track’, ca. 220 m, rainforest, ex |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
rotting log (R. W. Taylor); Hayfield near
Maprik ca. 150 m (R. W. Taylor); 13 km
NW Lae, Bubia, lowland rainforest (E. O.
Wilson); Finsch Harbor (L. Wagner); Maf-
fin Bay (E. S. Ross); Hollandia (H. A.
Levy); NW Japan I., SSE Sumberbaba,
Dawai R., jungle (H. Holtmann). Indone-
sia: Auraborine. Ambon (Mann); N. C. Ser-
am, Manusela N.P., Wae Mual Plain (M
J. D. Brendell); Seram, above Haruru,
near Masohi, 50 to 150 m, rainforest (W.
L. Brown); Irian Jaya, km 12 S of Sorong,
forest fragment (W. L. Brown); Irian Jaya,
Pi ereeport Concession, Siewa Camp.,
03.04°S, 136.38°E, 200 ft, lowland second
rainforest, #98-33 and #98-48 (on rotten
log), #98-63 (strays in leaf litter), #98-83
(under loose bark of log, Venom Voucher)
(R. R. Snelling). Micronesia: Ponape Is.,
Mt. Kubersoh, 1,900 ft (P. A. Adams);
Ponape Iss. 55 Tolotom, 1,700ut (P7A
Adams); Ponape Is., Mt. Dolennankap,
1,700 to 2,000 ft (H. K. Townes); Ponape
Is., near Colonia (Townes); Pohnpei, Mal-
en, above Kepirohi Falls, 350 m (Ron
Close); Pohnpei, Mall Island, in coconut
husk litter (Ron Clouse); Pohnpei, Quarter
Mile, upriver from Mahnd, nest in rotten
tree fern stump (Ron Clouse); Pohnpei,
above Kepirohi Falls, 350 m, both in rot-
ten log and on bracket fungus (Ron
Clouse).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter, in rotten branch, and under the
bark of a log.
Pristomyrmex quadridentatus (André)
Figures 163-166, 265, 275
Odontomyrmex quadridentatus André, 1905: 208.
Lectotype worker, designated by Taylor (1965: 43),
Australia: Sydney (Duchaussoy) (MNHN) (Taylor,
1965). [Note: Odontomyrmex quadridentatus An-
dré was automatically transferred to Pristomyrmex
quadridentatus (André), when Forel (1915: 53)
designated Odontomyrmex as a subgenus of Pris-
tomyrmex |.
Pristomyrmex (Odontomyrmex) quadridentatus var.
queenslandensis Forel, 1915: 53. Syntype workers,
Australia: Queensland, Mt. Tambourine, 1913
(Mjébergs) (MHNG) [examined]. [Synonymy by
Taylor, 1965].
463
Notes: (1) Although I did not examine
the lectotype of P. quadridentatus, 1 had
access to some specimens that have been
compared with it by R. W. Taylor. (2) A
lectotype and a paralectotype of P. quad-
ridentatus, designated by Taylor, belong to
self-colored golden-brown form (Taylor,
1965). But the three syntypes of P. quad-
ridentatus var. queenslandensis, seen by
myself, belong to the bicolored form, with
head, gaster, and appendages golden-
brown to reddish-brown but ARearale pet-
iole, and postpetiole dark-brown. (3) Tay-
lor (1965) mentioned that “judging from
subsequent records the Sydney type-local-
ity may be outside the true range of this
species and should be regarded with res-
ervation as a distributional record until
confirmed.”
Diagnosis ( Worker). Pronotal spines ca.
0.15 to 0.24, much longer and more robust
than propodeal armaments (ca. 0.06—0.09):
dorsal surfaces of head and alitrunk
smooth and shining, but a transverse ridge
present at the approximate position of me-
tanotal groove; petiole node and postpeti-
ole each with at least one pair of erect or
suberect hairs; ventral surface of clypeus
usually with a long transverse ridge.
Worker. TL < 3.344. AD, eb 0.86— lee
HW 0.80-1.08, Cl 93 =O SIE 0.86-1.16.
SI 97-113, EL 0.16—0.20, PW 0.52-0.67,
INE) OFS SSIs IW O22 0) 2X6). TRIP Ly Oe
0.28; PRI 93=109' Gr = 52).
Mandibles usually smooth and shining
but sometimes with a few longitudinal ru-
gae, varying from superficial to distinct.
Masticatory margin of mandible with three
teeth arranged as the strongest apical +
the second strongest preapical + a long
diastema + a truncated basal tooth. Basal
margin of mandible almost straight, lack-
ing a toothlike prominence. Clypeus with
a strong median longitudinal carina. An-
terior clypeal margin with three teeth: a
median tooth and one on each side. Ven-
tral surface of clypeus usually with a long
transverse ridge but sometimes showing
only two foothiike prominences. Palp for
mula 2,2. Frontal carinae usually extend-
464
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
BSS
Figures 163-166. Pristomyrmex quadridentatus (André). 163A: Worker head, full-face view; 163B: Showing a transverse ridge
on the ventral clypeus; 164: Worker, lateral view; 165A: Queen head, full-face view; 165B: Showing a transverse ridge on the
ventral clypeus; 166: Queen, lateral view.
ing to the level of the posterior margins of
eyes, but sometimes they are not so. Weak
scrobal impressions present lateral to the
frontal carinae. Frontal lobes weak: thus,
the antennal articulations are almost en-
tirely exposed. Antennal scapes, lying on
the dorsal head, slightly surpassing the oc-
cipital margin of head. Between the sec-
ond and seventh funicular segments of an-
tennae, the second is longest. Eyes usually
containing seven to eight ommatidia in the
longest row. Occipital carina distinct. Pro-
file shape of alitrunk and pedicel segments
as in Figure 164. Pronotal spines, ca. 0.15
to 0.24, much longer and more robust than
propodeal armaments, which are a pair of
teeth or short spines (ca. 0.06—0.09). Me- |
tapleural lobes usually elongate-triangular,
usually longer than propodeal armaments. —
Petiole node in profile, with the antero-
dorsal angle higher than the posterodorsal; |
in dorsal view, crest of the node convex.
Anterior and dorsal faces of the postpeti-
ole in profile forming a single curved sur-
face. Postpetiole in dorsal view broadening
from front to back. Dorsum of head
smooth and shining, except for a short ca-
rina present below the antennal scrobe,
subparallel to frontal carina. Dorsum of al- |
|
itrunk smooth and shining, with a distinct
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
transverse carina on the anterior margin of
the pronotum and a transverse ridge pre-
sent at the approximate position of metan-
otal groove. Petiole and _ postpetiole
smooth and shining. Gaster unsculptured.
Dorsal surfaces of head and alitrunk with
numerous erect or suberect hairs. Petiole
node and postpetiole each with bilaterally
distributed hairs, as shown in Figure 164.
First gastral tergite lacking erect or sub-
erect hairs. A few pairs of forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect to suberect short hairs. Body
self-colored (golden-brown) or bicolored
(i.e., head, gaster, and appendages golden-
brown to reddish-brown but alitrunk and
pedicel segments blackish-brown).
Queen. TL 3.56-5.02, HL 0.93-1.16,
HW 0.90-1.12, CI 93—L00, SL 0.93—1.12,
SI 96-103, PW 0.68—0.82, AL 1.02—1.30,
PPW 0.29-0.32, PPL 0.28—0.36, PPI S9-—
Hil (1 = 7).
General shape as in Figures 165-166,
with normal caste differences from the
conspecific worker; pronotum with a pair
of tubercles or minute teeth or sometimes
unarmed; propodeum with a pair of tu-
bercles or minute teeth; other characters
similar to worker.
Male (Figs. 265, 275). Four male spec-
imens are examined; two of them, collect-
ed in New South Wales (New England
Nat. Park, Platypus Creek, STRF) by
Nicholls, were originally mounted with
two and three self-colored workers, re-
spectively; one male, collected in SE
Queensland (Rathdowney, Lever’s Plateau,
Philp’s Farm) by J. B. Williams, and one
in Queensland (National Pk.), by F. A.
Perkins, were originally mounted with a
bicolored worker, respectively.
TL 3.63—4.14, HL 0.60—0.68, HW 0.62-—
0.72, CI 103-106, SL 0.22—0.26, SI 34—37,
HWE 0.84—0.93, EL 0.32—0.34, PW 0.82—
0.86, AL 1.22—-1.36, PPW 0.24—0.28, PPL
0.26—0.30, PPI 92-100 (n = 4).
Head, including the eyes, distinctly
broader than long. Clypeus convex, lacking
a median longitudinal carina, but with sev-
465
eral short rugae present on the posterior
clypeal margin. Anterior clypeal margin
straight. Palp formula 2,2; the second seg-
ments of maxillary and labial palps rather
long. Frontal carinae absent. Maximum di-
ameter of the median ocellus 0.08 to 0.10.
On the mesoscutum, notauli absent or
weakly impressed or fairly distinct; parap-
sidal furrows absent or superficially im-
pressed. Scuto-scutellar sulcus with 15 to
16 low, short ridges. Propodeum lacking
armaments. Metapleural lobes subtrian-
gular. Middle and hind tibiae without any
spurs. Petiole node in profile low and
rounded dorsally, with a long anterior pe-
duncle. Postpetiole low, in dorsal view usu-
ally longer than rarely as long as broad.
Posterior face of postpetiole almost entire-
ly attached to the first gastral segment.
Dorsum of head smooth and shining, but
frontal area usually with a median longi-
tudinal carina. Alitrunk smooth and shin-
ing, except for those marked sutures. Pet-
iole, postpetiole, and gaster smooth and
shining. All dorsal surfaces with abundant
long hairs. Scapes and tibiae with numer-
ous erect or suberect short hairs. Color
reddish-brown to blackish-brown; wings
infuscated.
Comments and Discussion. Pristomyr-
mex quadridentatus occurs in Australia. It
is closely related to P. wheeleri, also from
Australia. The workers of the two species
share the following critical characters: (1)
the masticatory margin of mandible with
three teeth arranged as an apical + a
preapical + a long diastema + a truncated
basal tooth; (2) palp formula 2,2; (3) the
anterior clypeal margin with three strong
teeth; (4) the clypeus with a median lon-
gitudinal carina; (5) the pronotum with a
pair of fairly long spines; (6) the dorsum
of alitrunk smooth and shining but with a
transverse carina present at the approxi-
mate position of metanotal groove; and (7)
first gastral tergite lacking erect or sub-
erect hairs. Pristomyrmex quadridentatus
differs from P. wheeleri in the workers and
queens, as follows:
466
P. quadridentatus
Propodeum with a pair of teeth or short
spines, ca. 0.06 to 0.09, much shorter
than pronotal spines (worker). Pro-
podeum tuberculate (queen)
Ventral face of clypeus with a long trans-
verse ridge or two toothlike promi-
nences (worker and queen)
Head relatively narrow (worker: CI 93—
101, HW 0.80-1.08; queen: CI 93-
100, HW 0.90-1.12)
Pronotum unarmed or with a pair of mi-
nute teeth (queen)
P. wheeleri
Propodeum with a pair of fairly long
spines, ca. 0.18 to 0.28, about equal
in length to or longer than pronotal
spines (worker); propodeum with a
pair of spines (queen)
Ventral center of clypeus usually with a
short transverse carina; sometimes,
this carina weak or indistinct (worker
and queen)
Head relatively broad (worker: CI 103—
116, HW 0.97-1.34; queen: CI 107—
AS IVVidl S242)
Pronotum armed with a pair of acute
short spines (queen)
Characters separating P. quadridentatus
from P. erythropygus, from the two Afri-
can species P. africanus and P. trogor, and
from the two Oriental species P. flavus and
P. collinus are provided under P. erythro-
pygus, under P. africanus, and under P. fla-
tus, respectively.
Taylor observes that there are two color
forms of P. quadridentatus. One is a uni-
form reddish-brown and occupies the
southern portion of its geographic range.
The other is bicolored (head, gaster, and
appendages reddish-brown but alitrunk,
petiole, and postpetiole dark-brown) and
occurs in more northern localities. Taylor
found no evidence of sympatry of the two
forms; thus, he postulated that the bicol-
ored form might be result of character dis-
placement where P. quadridentatus co-oc-
curs with P. wheeleri (Taylor, 1965, 1968).
However, six specimens, collected by P.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Ward in 1976, have the same records of
locality (New South Wales: Whian Whian,
SF, 28.39S/153.20E, rainforest, 200 m).
The six specimens consists of two workers
of each color form of P. quadridentatus
and two workers of P. wheeleri. This col-
lection implies that two forms of P. quad-
ridentatus partly overlap in their distribu-
tion (in other words, character displace-
ment is not the cause of the bicolored
form in areas of sympatry with P. whee-
leri). Of course, these data need to be fur-
ther confirmed because they come from
the labels on only a few specimens.
Taylor (1965) also proposed that the two
color forms “might prove ultimately to be
good biological _ species”, although there
are no Gites detectable morphological dif-
ferences between them. This question will
be clarified by further collecting: If the
two forms are never found to eoeust in the
same colony, they represent two good spe-
cies: otherwise, they belong to the same
species. (Note: To date, these two forms
have not been collected in the same col-
ony. )
The larva of P. quadridentatus was de-
scribed by Wheeler and Wheeler (1973).
Material Examined (ANIC, MCZC).
Self-colored form: Australia: New South
Wales: Grafton, Pt. lookout, rainforest (P.
F. Darlington); New England National
Park, Platypus Crk., STRF, in log (Nich-
olls); Dorrigo, 3,000 ft (W. Heron); Dor-
rigo NP, W rend Blackbutt Tr., subtropical
rainforest, 790 m (A. Nevton and M.
Thayer); East foot hills, Barrington Tops
near Cobark, rainforest, 400 ft (B. B. Low-
ery); Brunswick Heads, faunal reserve, RF,
sea level, under bark sheath of Bangalow
Palm (B. B. Lowery); Macksville, Warrell
Crk. area, RF valley, 150 ft, red rotten log
(B. B. Lowery); ca. 5 mi NW of Bruxner
Pk. (D. H. Colless); Upr. Allyn Val. near
Eccleston, rainforest, ca. 2,000 ft, ex rotten
log and nocturnal strays (Taylor and
Brooks): Whian Whian, SF, 28.39°S,
153.20°E, rainforest, 200 m, ca. 8 m above
ground, in epiphytic fern on tree, acc. no.
1721, 17.vi.1976 (PB. Ward); Coffs Harb
REVISION OF
Bruxner Pk., ca. 70 m (R. W. Taylor); Roy-
al Nat. Park, rainforest, 50 m, 34.09°S/
151.01°E, ex rotten log (P. Ward). Bicol-
ored form: Australia: Queensland: Mt.
D Aguilar Range, 2,000 ft, ex rotten log (R
W. Taylor); Stradbroke Is. (H. Hacker);
Tamborine Mt. (A. M. Lea); Tamborine
Mt., 1,800 ft (W. L. Brown); Tamborine
Mt., Cedar Creek, rotting logs (W. L.
Brown); Tamborine Mt., near Witches
Halls 27.07 5, 53. J0°E (R. J. Kohout);
Tamborine Mt., RF, 2,000 ft, rotten log (B
B. Lowery); 6 km SSW North Tamborine,
Pi O96 9/153°11’E, 500 m, #9828 (P S.
Ward); Binna Burra: 2,600 ft, rotten logs
(R. W. Taylor); Lamington N.P., Cocunenn
Gorge, ex rotten log (R. W. Taylor); Below
Eprinebrook Upper Tallebudgera Ck, RF,
550 m (Monteith, Thompson. and Cook);
Numinbah Arch, rainforest, 1,500 ft, ex
rotten log (B. B. Lowery); Joalah Nat.
Rank 27. 55°S, Sse cE CanooOnme (Rs VV
Taylor and R. Kohout); Rathdowney, Le-
vers Plateau, Philp’s Farm (J. B. Williams);
National Park (F. A. Perkins); Cooloola,
rainforest (P. J. M. Greenslade). New
South Wales: Wiangaree SF, Brindle Ck.,
740 m, subtropical rainforest (A. Newton
and M. Thayer); Mt. Warning, rainforest,
ex rotten log (B. B. Lowery); Mt. Warning,
rainforest, ca. 1,000 ft, under bark ice
of Bangalow Palm (B. B. Lowery); Nobby’s
Crk., 10 mi NW Murwillumbah, rainfo-
rest, ca. 1,500 ft, under bark sheath of
Bangalow Palm (B. B. Lowery); Whian
Whian SF, 28.39°S, 153.20°E, 200 m, acc.
no. 1709, 16.vi.1976 (P. Ward): Tomewin,
rainforest, 1,200 ft, on Trunk of fig, 8 ft off
ground, (B. B. Lowery); Acacia Plateau (J.
Armstrong); 14 km W = Urbenville, Too-
loom Plateau, 28.29°S/152.24°E (I. D.
Naumann).
Ecological Information. This species oc-
curs in rainforest. It forages nocturnally in
the open on logs, tree trunks, and low fo-
liage. Nests are located in rotting logs and
under the bark sheaths of Bangalow palms
(Taylor, 1965, 1968).
THE ANT GENUS PRISTOMYRMEX * Wang
467
Pristomyrmex quindentatus sp. n.
Figures 167-170
Diagnosis (Worker). Masticatory margin
of Honcible with five teeth; pronotum
with a pair of triangular teeth; dorsal sur-
faces of head and alitrunk with scattered
foveolate punctures.
Holotype Worker (MCZC). TL 3.54, HL
0.86, HW 0.84, CI 98, SL 0.79, SI 94, EL
0.12, PW 0.56, AL 0.83. Paratypes, five
workers and two queens (MCZC).
Worker ml, 311023545 Ee O81 0!86;
HW 0.79-0.84, CI 96—98, SL 0.76—0.80,
Si94S100. Hic On 1=0825 PW 10%53=0:56)
AL 0.78-0.86, PPW 0.22—0.23, PPL 0.21—
022PPI00>l0a"Ges— 16)!
Mandibles smooth and shining, except
for a few longitudinal rugae. Masticatory
margin of mandible with five teeth ar-
ranged as the strongest eee + the sec-
eine strongest preapical + a diastema +
three sell denticles of similar size (some-
times three small denticles are fused to-
gether so that they are not clearly visible;
or, one of them is weak or worn down, but
the length of diastema is slightly shorter
than the distance covered by these three
teeth). Basal margin of mandible lacking
toothlike prominences. Clypeus depressed
and smooth, but the median carina of the
frontal area extending a little to the clyp-
eus. Anterior clypeal margin usually with
a broad, truncated median denticle and
two to three others on each side. Ventral
center of clypeus with a prominent tooth.
Palp formula 1,3. Frontal carinae strong,
extending to the level of the posterior mar-
gins of eyes. Slightly impressed scrobal ar-
eas present lateral to the frontal carinae.
Frontal lobes weak; thus, the antennal ar-
ticulations are almost entirely exposed. An-
tennal scapes, laid on the dorsal head, just
surpassing the occipital margin of head.
Eyes moderately sized. Occipital margin
feebly concave. Pronotum with a pair of
triangular teeth. Propodeum with a pair of
short spines that are about two to three
times the length of pronotal armaments.
Metapleural lobes somewhat rounded.
468
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 167-170. Pristomyrmex quindentatus sp. n. 167A: Worker head, full-face view; 167B: Showing a tooth on the center
of ventral clypeus; 168: Worker, lateral view; 169A: Queen head, full-face view; 169B: Showing a tooth on the center of ventral
clypeus; 170: Queen, lateral view.
Petiole node in profile with a fairly long
anterior peduncle; its anterodorsal angle is
on a higher level than the posterodorsal
(Fig. 168). Postpetiole in profile rounded
dorsally, in dorsal view about quadrate.
Dorsum of head, except for the shallow
scrobes, with numerous scattered foveo-
late punctures. Dorsum of alitrunk pos-
sessing scattered foveolate punctures, usu-
ally with a smooth and unsculptured me-
dian strip. Petiole smooth and _ shining,
with a weak longitudinal ruga on each side.
Postpetiole and gaster unsculptured. Dor-
sal surfaces of head and alitrunk with nu-
merous erect or suberect short hairs. Two
pairs of hairs usually present on the dor-
sum of petiole node, and a few pairs on
the dorsum of postpetiole, as illustrated in
Figure 168. First gastral tergite lacking or
bearing few suberect hairs. A few pairs of
forward-projecting hairs present near the
anterior clypeal margin. Scapes and tibiae
with numerous erect to suberect short
hairs. Color reddish-brown. |
Queen. TL 3.64, 3.70; HL 0.83, 0.83; |
HW 0.82, 0.83; CI 99, 100; SL 0.72, 0.78; |
SI 88, 94; EL 0.18, 0.19; PW 0.64, 0.64; |
AL 0.97, 1.00: PPW 0.23, 0.23: PPI 0.228 |
O23 ERE LOOMO05a¢7— 2): |
Generally similar to worker, except for —
normal caste differences. In addition, |
pronotum unarmed; propodeal armaments
toothlike, shorter than those in the con-—
specific worker, and the dorsum of alitrunk |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
lacking an unsculptured median longitu-
dinal strip.
Male. Unknown.
Comments. Pristomyrmex quindentatus
is closely related to P. occultus. The two
species occur in the Oriental region. Their
workers share a critical character, that is,
the masticatory margin of mandible with
an apical tooth + a preapical tooth + a
diastema + three small denticles of similar
size; this dentition is not seen in all other
Pristomyrmex species, except in P. eduar-
di. But P. quindentatus and P. occultus can
be separated from P. eduardi in the work-
ers in having a pair of pronotal teeth.
The worker of P. quindentatus differs
from that of P. occultus as follows:
P. quindentatus
Dorsal surfaces of head and _alitrunk
only with scattered, shallow foveolate
punctures; dorsum of alitrunk with an
unsculptured median longitudinal
strip
Anterior clypeal margin with a truncat-
ed median tooth
A lateral carina lacking or vestigial on
each side of the petiole node
SL 0.76—0.80, SI 94—100
P. occultus
Dorsal surfaces of head and alitrunk en-
tirely covered with coarse rugoreti-
culum
Median tooth on the anterior clypeal
margin not truncated
Petiole node with a lateral longitudinal
carina on each side
SieO 72-0 os Sl o7—93
Pristomyrmex quindentatus may have
evolved from the ancestor of P. quadri-
dens. Pristomyrmex quindentatus can be
separated from P. quadridens as follows:
the masticatory margin of the mandible
possesses five teeth in the workers and
queens of P. quindentatus but at most four
teeth in P. quadridens; in addition, the
propodeum is armed with a pair of short
spines in the workers of P. quindentatus
but a pair of teeth in P. quadridens.
Holotype Worker. Indonesia: Seram,
469
above Haruru, near Masohi, rainforest, 50
to 150 m, 18.iii. 1981 (W. L. Brown).
Paratypes. Five workers and two queens
with same data as holotype.
Additional Material Examined (MCZC).
A worker collected in Indonesia (Blawan,
Idjen, Dammerman, 950 m) has the fol-
lowing measurements: TL 3.50, HL 0.97,
HW 0.95, CI 99, SL 0.84, SI 88, EL 0.14,
PW 0.62, AL 0.94. It shows some varia-
tion: (1) The propodeum is armed with a
pair of triangular teeth, (2) the dorsum of
the head possesses dense foveolate punc-
tures, (3) a longitudinal carina is present
on each side of the petiole, and (4) three
small denticles on the masticatory margin
of the mandible are worn down and are
not clearly visible.
Ecological Information. This species has
been collected in rainforest.
Pristomyrmex sulcatus Emery stat. n.
Figures 171-176, 266, 276
Pristomyrmex brevispinosus sulcatus Emery, 1895:
464. Syntype workers, queen, Burma: Carin Cheba,
500 to 1,000 m, xii.1887 (L. Fea) [syntype workers
(MCSN, NHMYV) examined].
Diagnosis (Worker). Ventral surface of
clypeus with a strongly prominent tooth at
the center; pronotum with a pair of mod-
erately long spines (0.14—0.20), usually
longer than propodeal armaments (0.07—
0.13); dorsum of head with foveolate-retic-
ulate sculpture or rugoreticulum; petiole
node lacking foveolate punctures; first gas-
tral tergite lacking erect or suberect hairs.
Worker. TL 3.98—4.74, HL 0.98-1.16,
HW 0.96—-1.10, CI 93-105, SL 1.02—1.15,
SI 101—11t, El 0,140.19, Pw 0:64-0:75,
AL 0.98—1.18, PPW 0.26—0.30, PPL 0.26—
0.32, PPI 90-100 (n = 30).
Mandibles usually with some longitudi-
nal rugae, varying from superficial to dis-
tinct. Masticatory margin of mandible with
four teeth arranged as the strongest apical
+ the second strongest preapical + a long
diastema + two small basal teeth of similar
size that are fused together. Basal margin
of mandible almost straight, lacking a dis-
tinctly convex lobe or tooth. Clypeus with
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
A470
Aas
ie
“es
oF
—176. Pristomyrmex sulcatus Emery. 171A: Syntype worker head, full-face view; 172: Syntype worker, lateral view; |
173A: Non-type worker head, full-face view; 174: Non-type worker, lateral view; 175A: Queen head, full-face view; 176: Queen, |
lateral view; 171B, 173B and 175B: Showing that a prominent tooth is present on the ventral center of the clypeus in the syntype |
worker, non-type worker and queen, respectively.
Figures 171
|
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
a median longitudinal carina. Anterior
clypeal margin with a median denticle and
two to three others on each side; but
sometimes two lateral denticles are fused
into a larger tooth. Ventral center of clyp-
eus with a strongly prominent tooth. Palp
formula 1,3. Frontal carinae strong, ex-
tending to the level of the posterior mar-
gins of eyes, forming the dorsal margins of
the shallow antennal scrobes. Frontal
lobes very weak; thus, the antennal artic-
ulations are almost entirely exposed. An-
tennal scapes, when lying on the dorsal
head, slightly surpassing the occipital mar-
gin. Eyes usually containing six to seven
Bemacdia in the longest row. Profile
shape of alitrunk and pedicel segments as
in Figures 172 and 174. Pronotmn with a
pair of spines, varying in length from 0.14
(in the type series) to 0.20. Propodeum
with a pair of teeth or short spines varying
from 0.07 to 0.13. Metapleural lobes sub-
triangular or somewhat rounded. Petiole
Ride. in profile high, with a fairly long an-
terior peduncle; its anterodorsal angle is
on a higher level than the posterodor sal.
Postpetiole in profile rounded dorsally, in
dorsal view broadening from front to back,
about as long as or slightly longer than
broad. Dorsum of head, except for the
scrobal areas where rugae are somewhat
weak, with course rugoreticulum or dense
alveolate punctures. samulae but slightly
sparse sculpture present on the dorsum of
alittunk and often on the two sides of
pronotum. Petiole, postpetiole, and gaster
smooth and _ shining.
head and alitrunk <a numerous erect or
suberect hairs. Usually, two pairs of hairs
present on the dorsum of petiole node and
one to two pairs on the dorsum of post-
petiole. First gastral tergite lacking erect
or suberect hairs. A few pairs of forward-
projecting hairs present near the anterior
clypeal margin. Scapes and _ tibiae with
some erect to suberect hairs. Color red-
dish-brown.
Queen. TL 4.26-5.00, HL 0.98-1.10,
PIV, 1 OO=114- Cl 102106) SEARO 1. 12:
Hes 101 LE 012220264 PW.0!77_-0.90:
Dorsal surfaces of
471
AL 1.16—1.30, PPW 0.28—0.32, PPL 0.29-—
0.32, PPI 97-100 (@ = 4)
General shape as in Figures 175-176,
with normal caste differences from the
conspecific worker; pronotum usually un-
armed but rarely with a pair of minute
spines. Other characters similar to worker.
The queens of P. sulcatus are almost in-
distinguishable from those of P. brevispi-
nosus at present.
Male (Figs. 266, 276). One male speci-
men (MCZC), collected in Thailand (Nak-
hon Ratchasima Prov., Khao Yai Nat. Park,
750 m, hill forest) by I. Burikam and W.
L. Brown, was originally mounted with a
worker on the same pin; it has the follow-
ing measurements: TL 3.90, HL 0.58, HW
OOL GET OSs SiO 22S Son ELV OsS35
EE 0733) 5 PW 01S2,Ale 1elsePRW 10:24"
PRPs 0:24> PRP l00;Ga—s))
Head, including the eyes, distinctly
broader than long. Clypeus convex, with-
out a median longitudinal carina. Palp for-
mula 1,3. Frontal carinae weak and short,
just reaching the level of the posterior
margins of antennal insertions. Maximum
diameter of the median ocellus 0.10. On
the mesonotum, notauli strongly marked,
forming a Y shape; parapsidal AunGuis ab-
sent. Salo: scutellar sulcus rather broad,
with 10 ridges that expand at the upper
end. Propodeum slightly tuberculate, lack-
ing teeth or spines. Metapleural lobes sub-
tr iangular. Middle and hind tibiae without
any spurs. Petiole node in profile low, with
a fairly long anterior peduncle. Postpetiole
low, in dorsal view about as long as broad.
Dorsum of head generally oot and
shining but with a median longitudinal ca-
rina present on the frontal area fand several
short rugae on the posterior clypeal mar-
gin. Alieeani smooth and shining, except
for those marked sutures. Petiole, post-
petiole, and gaster smooth and shining. All
dorsal surfaces with abundant erect or
suberect hairs. Scapes and legs with erect
or suberect short hairs. Body reddish-
brown; funicular segments of antennae
white and wings slightly light-yellow. At
the present, dhe male of P. Ruleatis is al-
most indistinguished from the males of P.
brevispinosus and P. quadridens.
Discussion and Comments. Taxonomic
status of “sulcatus” is somewhat compli-
cated. The syntype workers of P. brevispi-
nosus sulcatus differ from those of P. bre-
vispinosus by possessing a pair of fairly
long pronotal spines (0.15—0.16), as com-
pared with a pair of teeth (0.06) in the
latter. However, after examining all avail-
able material, I find that the length of the
pronotal armaments is continuously vari-
able (from 0.06, as in the syntypes of P.
brevispinosus, to 0.20, as in the specimens
from Khao Yai Nat. Park, Thailand). The
syntype workers of “sulcatus”, in fact, are
intermediates between the two extreme
ends (i.e., in one extreme, the pronotal
teeth are slightly shorter than or about as
long as the propodeal teeth; in the other
extreme, the pronotal spines are two to
three times as long as the propodeal ar-
maments; see Figs. 114, 172, 14D), Te keep
“sulcatus” as a valid name (i.e., raise it to
the rank of species instead of assigning it
as a junior synonym of P. brevispinosus)
because more ecological work must be
done before the status of “sulcatus” be-
comes clarified. With this tentative pro-
posal, P. sulcatus comprises those popula-
tions with pronotal spines (ca. 0.14—0.20),
distributed in Pahang (1,300-1,720 m),
Malaya, and in northwest and central Thai-
land, Burma, Nepal, and southwest China.
Pristomyrmex brevispinosus comprises
populations with toothlike pronotal arma-
ments (ca. 0.06—0.10), occurring in Pahang
(1,250 m), Malaya, and Trang Province of
South Thailand (07.55°N) and in the Phil-
ippines, Taiwan, and Japan. In other
words, P. sillectans may be a northerly re-
placement OEE. brevispinosus along the
Malay peninsula.
The separation of P. sulcatus from P.
costatus is provided under P. costatus. The
worker of P. sulcatus differs from that of
P. hirsutus by its petiole node with the an-
terodorsal angle higher than the postero-
dorsal and its first gastral tergite lacking
erect or suberect hairs. The worker of P
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
f
sulcatus differs from that of P. modestus
by its petiole node higher than long and |
lacking distinct foveolate punctures. The |
worker of P. sulcatus differs from that of |
P. occultus by its masticatory margin of the |
mandible possessing at most four teeth |
and its propodeal armaments not longer
than the pronotal spines. The queen of P.
sulcatus differs from that of P. bicolor by.
possessing a strongly prominent tooth on |
the ventral center of the clypeus. The |
worker and queen of P. sulcatus differ |
from those of the Australian P. foveolatus
and P. thoracicus as follows: |
P. sulcatus
Pronotal spines longer than or about as_
long as the propodeal armaments |
(worker) |
Propodeum with a pair of teeth (Fig.
176) (queen)
Maxillary palp with one segment (work-_
er and queen) |
P. foveolatus and P. thoracicus
Pronotal spines much shorter than the
propodeal spines (worker)
Propodeum with a pair of fairly long
spines (Figs. 143, 180) (queen)
Maxillary palp with two segments (work-_
er and queen)
Material Examined (ANIC, BMNH, |
IZAS, LACM, MCZC, MHNG, NHMV,}
USNM). Thailand: Nakhon Ratchasima
Prov., Khao Yai Nat. Park, 700 to 750 m,|
hill forest, rotten wood (I. Burikam and W. |
L. Brown); Khao Yai Nat. Park (L6bl and!
Burckhardt); Chiang Mai (54 km NE
Chiang Mai), Mae Nang Kaeo, 900 m)
(Burckhardt and L6bl); Chiang Mai Pr. st
Doi Suthep NP (7), near Ruesse Care, 900 |
to 1,000 m (Zettel); Chiang Mai Prov., |
Chiang Dao Wildlife Sanctuary, 19. 29°N |
98. 97°, 470 to 500 m elevation (Saowana_
Sontintclnet). Nepal: Khandbari District,
Arun River at Num 1,500 to 1,600 m (A. /
and Z. Smetana). Burma: Carin Cheba (L. 7
Fea). China, Yunnan Prov. (?). W. Malay- |
sia: Pahang, Cameron Highls, Gunung Ja-_
sar, 1,720 m (L6blI and Calame); Malaysia:
Pahang, Cameron Highlands, Tanah Rata |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
473
Figures 177-180. Pristomyrmex thoracicus Taylor. 177A: Worker head, full-face view; 177B: Showing a tooth on the center of
ventral clypeus; 178: Worker, lateral view; 179A: Queen head, full-face view; 179B: Showing a toothlike prominence on the
center of ventral clypeus; 180: Queen, lateral view.
Umg., Gn. Jasar, 1300 m (Schuh and
Lang)?: Kabu, 400 ft (Abor Exped.).
Ecological Information. This species has
been collected in forests.
Pristomyrmex thoracicus Taylor
Figures 177-180
Pristomyrmex thoracicus Taylor, 1965: 41. Holotype
worker, Australia: N. Queensland, Lake Eacham
National Park, rainforest, 2,500 ft, 6.vi.1962 (R. W.
Taylor) (MCZC) [examined].
Diagnosis (Worker). Pronotum with a
pair of triangular short spines (ca. 0.06—
0.08); propodeal spines long (ca. 0.19-
0.24); dorsum of head, except for the an-
tennal scrobes, with foveolate-reticulate
sculpture; postpetiole unsculptured; PPI
109-121; SL 0.86—0.98 and SI 97-103.
Worker. TL 3.22-3.72, HL 0.86—0.96,
HW 0.86-—0.96, CI 97-101, SL 0.86—0.98,
SI 97-103, EL 0.10—0.13, PW 0.54—0.60,
AL 0.84—0.96, PPW 0.23—0.26, PPL 0.20—
O22 SE RIGhOS= 24)
Mandibles with a few longitudinal ru-
gae. Masticatory margin of mandible with
three teeth arranged as an apical + a
preapical + a long diastema + a truncated
basal tooth. Basal margin of mandible lack-
ing a distinct curved lobe or tooth. Clyp-
eus with a strong median longitudinal ca-
rina. Anterior clypeal margin with a me-
dian denticle and usually two others on
A474
each side, but sometimes two lateral den-
ticles are fused into a larger tooth. Ventral
center of clypeus with a low, broad, tooth-
like prominence. Palp formula 2,3 . Frontal
carinae well developed, beyond i level
of the posterior margins of the eyes. Scro-
bal areas shallow, present lateral to the
frontal carinae in full-face view. Frontal
lobes almost completely absent so that the
antennal articulations are entirely exposed.
Antennal scapes usually slightly surpassing
the occipital margin of head when lying in
the antennal scrobes. Eyes with five to six
ommatidia in the longest row. Profile
shape of alitrunk and pedicel segments as
in Figure 178. Pronotum with a pair of tri-
angular short spines, ca. 0.06 to 0.08. Pro-
podeal spines long, ca. 0.19 to 0.24, usually
slightly upeurved at their apices. Meta-
pleural lobe small-triangular with a rather
acute apex. Petiole node in profile higher
than long, with a long anterior peduncle,
its adicrodoteal angle forming an apex and
its dorsum sloping dower posteriorly.
In dorsal view, crest of petiole node
strongly convex. Postpetiole in profile
rounded dorsally, in dorsal view broader
than long and broadening from front to
back. Dorsum of head between the frontal
carinae, as well as the two sides of the dor-
sal head, with foveolate-reticulate sculp-
ture. Antennal scrobes rather smooth, with
only a few weak rugae. Dorsum of anten-
nal scape with a longitudinal carina. Dor-
sum of alitrunk with a rugoreticulum. Pet-
iole and postpetiole smooth and _ shining.
Gaster unsculptured. Dorsal surfaces of
head and alitrunk with numerous erect or
suberect hairs. Dorsal surfaces of petiole
node and postpetiole each with a pair of
hairs. First gastral tergite lacking erect or
suberect fires A few pairs of fee pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect to suberect short hairs. Color
reddish-brown.
Queen. TL 3.844.10, HL 0.94—0.96,
HW 0.94—0.96, CI LOO—100, SL 0.94—0.96,
SI 98-100, EL 0.16—0.18, PW 0.70_-0.72,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Al e061 4S PPWa0.25=0)27 SE Rin 022=
O225P RIAA 23h os
General shape as in Figures 179-180,
with normal caste differences from the
conspecific worker; pronotum unarmed;
crest of petiole node in dorsal view feebly |
convex; other characters similar to worker. |
Male. Unknown. |
Comments. Pristomyrmex thoracicus is |
so far known only from North Queensland, |
Australia. It is a sibling species of P. fov- |
eolatus, also from North Queensland. |
Their differences are given under P. fov-
eolatus. Characters separating P. thoraci- |
cus from the Asian P. brevispinosus, and
from the African P. cribrarius (a member -
of the cribrarius group) are provided un-
der P. foveolatus and under P. cribrarius, |
respectively. |
Material Examined (MCZC, ANIC). |
Australia: Queensland, Lake Eacham, -
rainforest, 2,500 ft, nest ex rotten log (R. |
W. Taylor); N.Q., Kuranda, Tainiorest |
1,100 ft, stray floor (R. W. Taylor); N.Q., |
Cravtonds Lookout, Beatrice River (Dar-.
lingtons); N.Q., Malanda, rainforest, rotten |
log (W. L. Brown); N.Q., 3.2 km E of Lake
Barrine, ca. 700 m, rainforest, ex rotten.
log (R. W. Taylor and J. Feehan); N.Q.,}
Iralce Barrine Nat. Pk., 760 m, 17.l5°Sm
148.38°E, rainforest, ex rotting log (R. W.
Taylor and T. A. Weir); N.Q., Palmerston |
N.P.. ca. 1,000 ft, rainforest, nest in soil |
under log (R. W. Taylor). |
Ecological Information. This species oc- |
curs in rainforest nesting in rotting logs.
and in soil under logs (Taylor, 1965, 1968). |
|
Pristomyrmex trachylissus (F. Smith)
Figures 181-184
Myrmica trachylissa F. Smith, 1858: 126. Holotype |
queen, Borneo (A. R. Wallace) (BMNH) [exam- |
ined].
Pristomyrmex trachylissus (F. Smith) Mayr, 1886: |
359.
Diagnosis (Worker). Large size (HL = |
1.36, HW Ses): masticatory margin of
mandible with five teeth; pronotum Heh a |
pair of exceptionally long spines that are |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
OS
J
Sy
ey,
L 3
ZEN
A475
Figures 181-184. Pristomyrmex trachylissus (F. Smith). 181A: Worker head, full-face view; 181B: Showing a short ruga on the
ventral clypeus; 182: Worker, lateral view; 183A: Queen head, full-face view, antennae missing; 183B: Showing a short ruga on
the ventral clypeus; 184: Queen, lateral view.
about as long as the distance between their
bases.
Worker. TL 5.58-6.48, HL 1.36-1.46,
IVY 1-45—1.62, Cl 107-114, SL 1.40=1.50,
ol 90-98, EL 0.22-0.24 PW 0.92-1.02,
AL 1.50-1.64, PPW 0.35—0.39, PPL 0.44—
0.48, PPI 76-83 (n = 6).
Mandibles generally smooth and shin-
ing, except for a few superficial rugae.
Masticatory margin of mandible with five
teeth arranged as the strongest apical +
the second strongest preapical + a small
third tooth + a short diastema (or this di-
astema indistinct) + two small basal teeth.
Basal margin of mandible with a central,
broadly curved lobe. Clypeus with a strong
median longitudinal carina. Anterior clyp-
eal margin with a median denticle and a
broad prominence on each side. Ventral
surface of clypeus generally smooth and
shining but sometimes with a weak, short,
transverse ruga at the center. Frontal ca-
rinae strong, extending to the level of the
posterior margins of the eyes. Antennal
scrobes shallow, present lateral to the fron-
tal carinae. Frontal lobes weak; thus, the
antennal articulations are almost entirely
exposed. Antennal scapes, laid on the dor-
sal head, slightly surpassing the occipital
margin of head. Eyes with 10 to 11 om-
matidia in the longest row. Profile shape of
alitrunk and pedicel segments as in Figure
182. Pronotum armed with a pair of ro-
bust, exceptionally long spines, ca. 0.40 to
0.50, about as long as the distance be-
tween their bases. Propodeum with a pair
of short spines, ca. 0.12 to 0.18, shorter
than or at most about as long as the dis-
476
tance between the bases of two propodeal
spines. Metapleural lobes each with a
somewhat blunt-rounded apex. Petiole
node in profile high, with a long anterior
peduncle; its anterodorsal angle is on a
higher level than the posterodorsal. Post-
petiole in profile rounded dorsally, in dor-
sal view longer than broad and broadening
from front to back. Dorsum of head en-
tirely sculptured with well-developed
coarse rugoreticulum. Similar sculpture
present on the dorsum of alitrunk and on
the two sides of pronotum, except for the
space between the bases of two pronotal
spines, which is rather smooth. Petiole
smooth and shining but with a lateral lon-
gitudinal carina on each side. Postpetiole
and gaster unsculptured, smooth, and
shining. Dorsal surfaces of head arial ali-
trunk moth numerous erect or suberect
hairs. A few pairs of hairs present on the
dorsum of petiole node and at least a pair
on the postpetiole, as shown in Figure 182.
First gastral tergite lacking ATE or sub-
erect “Thetin, A ian pairs ae forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect or suberect hairs. Color red-
dish-brown, but the masticatory margins
and the basal margins of mandibles and
the funiculi of antennae black-brown.
Queen. TL 7.55, HL 1.84, HW 2.14, CI
116, SL- (antennal scapes missing), SI-,
PW 52) Ale 2:10) (2 = 1):
General shape as in Figures 183-184,
with normal caste differences from the
conspecific worker; pronotum unarmed;
other characters similar to worker.
Male. Unknown.
Comments. Pristomyrmex trachylissus
must have evolved from the ancestor of P.
bicolor. It is extremely similar in appear-
ance of the workers and queens to P. bi-
color. The two species may occur sympat-
rically in Sarawak and in Sabah. Their
workers and queens can be separated by
the following characters:
P. trachylissus
Masticatory margin of mandible with
five teeth; diastema indistinct or very
short
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Basal margin of mandible with a central,
broadly curved lobe
Anterior clypeal margin with a median
tooth and a broad, short pone
on each side
SI < 100 (only in worker).
P. bicolor
Masticatory margin of mandible with.
four teeth; a long diastema present
between the preapical and the third
tooth
Basal margin of mandible lacking a dis-
tinctly curved lobe
Anterior clypeal margin usually with
seven denticles, but in some speci-
mens, two or three lateral denticles
fused into a broadly convex lobe
SI > 105 (only in worker).
Material Examined (BMNH, MCZC,
ANIC). East Malaysia: Sarawak, 4th Div.,
G. Mulu Nat. Pk., RGS Expd., Long pala,
lowland Pintorese on log and on rotten
log (B. Bolton); North Borneo (SE), For-
est Camp, 19 km N of Kalabakan 180 m
(Y. Hirashima).
Ecological Information. This species oc-
curs in rainforest and has been collected
on a rotten log.
Pristomyrmex trogor Bolton
Figures 185-186
Pristomyrmex trogor Bolton, 1981: 287. Holotyped
worker, Zaire (B. Congo on the label): S slope of.
Mt. Kahuzi, 1,900 m, 5.ix.1957 (E. S. Ross and R.
E. Leech) [CASC (Bolton, 1981)]; [five paratype |
workers (MCZC) examined].
Diagnosis (Worker). Frontal carinae ab--
sent; ventral surface of clypeus with two.
toothlike prominences; pronotum with a
pair of short spines; dorsal surfaces of head |
and alitrunk smooth and shining; petiole
and postpetiole lacking hairs.
Worker. TL 3.42-— 3.98, HL 0.88-1. 00,
HW 0.92-1.02, CI 102-105, SL 0.96—1.06, |
SI 100-104, EL 0.14—0.16, PW 0.56—0.62, |
AL 0.84—0.94, PPW 0.23-0.26, PPL 0.22—|
OLR6SRPI100=106KGis—ao):
Mandibles smooth, with only a few weak.
longitudinal basal rugae. Masticatory mar-_
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang AT
~l
Figures 185-186. Pristomyrmex trogor Bolton. 185A: Worker head, full-face view; 185B: Showing two toothlike prominences
on the ventral clypeus; 186: Worker, lateral view.
gin of mandible possessing the strongest
apical tooth + the second strongest preap-
ical tooth + a diastema + two small basal
denticles that are often fused into a broad,
short tooth. Basal margin of mandible
lacking a toothlike prominence or curved
lobe. Clypeus lacking a median longitudi-
nal carina. Ventral surface of clypeus with
two strongly prominent teeth. Anterior
clypeal margin with a median tooth and
two lateral denticles on each side; some-
times two small lateral denticles are fused
into one prominence. Palp formula 2,3
(Bolton, 1981). Frontal carinae absent.
Antennal scrobes absent. Frontal lobes in-
distinct. Antennal scapes, when lying on
the dorsal head, just reaching or slightly
surpassing the occipital margin of head.
Eyes containing five to six ommatidia in
the longest row. Promesonotum in dorsal
view weakly concave. Pronotum and pro-
podeum each with a pair of short spines
(Fig. 186). Metapleural lobes rounded. In
profile view, petiole node high, higher than
long, with a long anterior peduncle, its an-
terodorsal angle higher than the postero-
dorsal. In dorsal view, petiole node about
as broad as long. Postpetiole in profile
higher than long, rounded dorsally, in dor-
sal view broadening from front to back. All
dorsal surfaces unsculptured, smooth, and
shining. Dorsum of head with some fine
long hairs. Dorsal surface of alitrunk with
only one to two pairs of hairs that arise
from the lateral margins of the mesono-
tum. Petiole node, postpetiole, and first
gastral tergite lacking erect or suberect
hairs. A row of fine, forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with some fine,
appressed or decumbent, or suberect
hairs. Color reddish-brown.
Queen. Unknown.
Male. One paratype male has the same
data as holotype and was originally mount-
ed together with two paratype workers on
the same pin: TL 2.92) HL 0155, HW 0.57,
CHAl04 SEO S05 a3. FIVE O94 Re
0:28, PW. 0:82; Aly 16 @ = 1):
Head, including the eyes, distinctly
broader than long. Clypeus narrow and
convex, its anterior margin rather straight.
Frontal carinae absent. Maximum diame-
ter of the median ocellus 0.10. Scapes lon-
ger than the other antennal segments, ex-
cept for the apical ones. On the meso-
scutum, notauli distinct, forming a Y
shape; parapsidal furrows absent. Scuto-
scutellar sulcus with about 10 narrow short
ridges. Propodeum weakly tuberculate,
lacking teeth or spines. Metapleural lobes
subtriangular. Middle and hind tibiae with-
out any spurs. Waist abnormally shaped,
with a huge segment formed by the fusion
478
of petiole and postpetiole; entire posterior
face of the waist attached to first gastral
segment. Dorsum of head smooth and
shining, but frontal area with a median
longitudinal carina; a short ruga present
below each antennal socket. Alitrunk gen-
erally smooth and shining, except for those
marked sutures. Petiole, postpetiole, and
gaster smooth and shining. All dorsal sur-
faces with abundant long haus. Scapes and
tibiae with numerous erect or suberect
short hairs. Color somewhat blackish-
brown; wings white.
Con menis Pristomyrmex trogor is sim-
ilar to another African species, P african-
us, in many characters of the workers, such
as dentition of the masticatory margin of
mandible; two teeth present on the ventral
clypeus; promesonotum in dorsal view
shallowly concave or flat; pronotum and
propodeum each with a pair of short
spines; the structure and shape of petiole,
postpetiole, and metapleural lobes; as well
as the distribution of hairs. But P. trogor
is easily distinguished from P. africanus
and from the other African species of the
genus because P. trogor is the only species
lacking frontal carinae in the workers. In
Added. the dorsum of the head is smooth
and shining in the workers of P. trogor but
has foveolate punctures in P. africanus, P.
fossulatus, and P. cribrarius; the pronotum
possesses a pair of short spines in the
workers of P. trogor that is not seen in P.
fossulatus and P. orbiceps.
The separation of P. trogor from the two
Asian species (P. flatus and P. collinus) and
from the three Australian species (P. whee-
leri, P. erythropygus, and P. quadridenta-
tus) is given under P. flatus and under P.
africanus, respectively.
Distribution. Zaire (known only from
the type series).
Ecological Information. Unknown.
Pristomyrmex wheeleri Taylor
Figures 187-192
Pristomyrmex wheeleri Taylor, 1965: 48. Holotype
worker, Australia: SE Queensland, ca. Binna Burra,
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 |
rainforest, 2,800 to 3,000 ft, nest under stone, 21—|
25.v.1962 (R. W. Taylor) (MCZC) [examined].
i
Diagnosis (Worker). Masticatory margin |
of mandible with three teeth; antenet™ :
clypeal margin at most with three teeth;
propodeal spine length ca. 0.18 to 0.28,)
about equal to or slightly longer than pro-|
notal spines; dorsum of alitrunk smooth,
but with a transverse ridge present at the
approximate position of Perot groove;
petiole node and postpetiole dor sally witht
some hairs; first gastral tergite lee
hairs.
Worker, Wii 326-454. Hi 0.92-1" 17,
HW 0.97—1.34, Cl 103-116, SL 0:94=1 iss |
SI 88-100, EL 0.17—0.24; PW 0.56-0.74, |
AL 0.88—1.08, PPW 0.24—0.31, PPL 0.22—
0.28, PPI 104-122 (7 = 40).
Mandibles usually smooth and shining,
except for some small hair pits. Mastica-
tory margin of mandible with three teeth
arranged as an apical + a preapical + a|
long ate + a somewhat truncated
Racal tooth. Basal margin of mandible lack-
ing a curved lobe or roan. Clypeus with a
Lace longitudinal carina. Anterior clyp-.
eal margin usually with three strong teeth:
a Sateen tooth and one on each side, but
sometimes the median tooth weak or ves-.
tigial. Ventral surface of clypeus usually
with a short transverse carina. Palp for-
mula 2,2. Frontal carinae not or just ex-|
tending to the level of the posterior mar-
gins of the eyes. Antennal scrobes absent.
Frontal lobes nearly completely absent;
thus, the antennal articulations are almost)
entirely exposed. Antennal scapes, laid on
the dorsal head, slightly surpassing the oc-|
cipital margin. Hye. usually containing:
eight to nine ommatidia in the longest row. |
Profile shape of alitrunk and pedicel seg-
ments as in Figure 188. Pronotal spines:
varying in length, from 0.12 to 0.28. Pro-|
podeal spines Ree 0.18—0.28) equal to or}
slightly longer than the pronotal spines. |
Metapleural lobes small, usually triangular.
Petiole node in profile cath a long anterior.
peduncle; its anterodorsal angle is on a
higher level than the posterodor sal: some-|
|
|
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang A479
wie
; \
a
Figures 187-192. Pristomyrmex wheeleri Taylor. 187A: Worker head, full-face view; 187B: Showing a short ruga on the ventral
clypeus; 188: Worker, lateral view; 189A: Ergatoid queen, full-face view; 189B: Showing a short ruga on the ventral clypeus;:
190: Ergatoid queen, lateral view; 191A: Queen head, full-face view; 191B: Showing a short ruga on the ventral clypeus; 192:
Queen, lateral view.
480
times posterodorsal angle indistinct, show-
ing a single curved surface. Postpetiole in
profile as in Figure 188, in dorsal view
slightly broader than long and broadening
from front to back. Dorsum of head
smooth and shining. Dorsum of alitrunk
smooth, but with a transverse ridge pre-
sent at the approximate position of metan-
otal groove. Petiole, postpetiole, and gaster
smooth and shining. Dorsal surfaces of
head and alitrunk with numerous erect or
suberect long hairs. A pair of long hairs
bilaterally distributed on the dorsum of
petiole node and on the postpetiole, re-
spectively; sometimes the crests of both
petiole node and postpetiole with addi-
tional one to two pairs of short hairs. First
gastral tergite lacking erect or suberect
ours. Usually, three pairs of forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect to suberect short hairs. Color:
head deep reddish-brown; alitrunk, pedi-
cel segments, and gaster yellow-brown to
reddish-brown.
Ergatoid Queen. TL 4.06, HL 1.08, HW
WANG 1 WOW SE Le SIMS TRIE OPAL EY
OOSEPAL, 106, PST (0584 BSE2 0:25:
RPWi80:28)) PPPs 0265 PEN 108 @ =o).
Note: This is a paratype).
General shape as in Figures 189-190.
Similar to worker; color and pilosity as in
worker, but head with three ocelli and pro-
mesonotum convex. Flight sclerites and
wings lacking, but a black speck is present
on each lateral margin of the mesonotum.
Queen. TL 4.62—5.06, HL 1.09-1.28,
EValelS Ae ClO ald Silzel-O06=122.0)
SI 82-90, EL 0.22-0.28, PW 0.80-0.90,
AL 1.14-1.34, PPW 0.30-0.36, PPL 0.26—
OSOSBPl lO s3G — to)!
General shape as in Figures 191-192,
with normal caste differences from the
conspecific worker; a pair of acute minute
spines present on the humeral angles of
the pronotum; propodeal spines rather
long, ca. 0.20 to 0.24; other characters sim-
ilar to worker.
Male. Unknown.
Comments. Pristomyrmex wheeleri oc-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
curs in East Australia. Its two close rela-
tives are P. erythropygus and P. quadri-
dentatus, also from East Australia. Char-
acters separating P. wheeleri from P. ery-
thropygus and from P. quadridentatus are
provided under P. erythropygus and under
P. quadridentatus, respectively. The differ-
ences between P. wheeleri and the two
Asian species (P. flatus and P. collinus) and
between P. wheeleri and the two African
species (P. africanus and P. trogor) are giv-
en under P. flatus and P. africanus, re-
spectively. |
Material Examined (ANIC, MCZC).
Australia: SE Queensland: Mt. D’Aguilar.
Range, 2,000 ft, rainforest, ex rotten log
(R. W. Taylor); Cunningham's Gap, 3,000
ft, rainforest floor, ex small wood fragment
(R. W. Taylor); Cunningham’s Gap, rain-
forest, 2,500 ft, nest under stone (B. B..
Lowery); National Pk. (H. Hacker); Tam-
borine Mt., rotting leaves (A. M. Lea);;
Tamborine Mt., S side, Curris Falls, rain-.
forest, berlesate leaf mold (T. E. Wood-
ward); Tamborine Mt., rainforest, 2,000 ft,
nest between stones (B. B. Lowery); ca.
Binna Burra, rainforest, 2,600 to 3,000:
ft, nest under stone (R. W. Taylor);
M’Pheraon Rge, v. Binna Burra, rainforest,
2,600 to 3,600 ft (P. F. Darlington); Binna’
Burra, Lamington Nat. Pk., leaf and log
litter (J. and N. Lawrence); Lamington|
Nat. Park (O’Reillys), 28.14°S, 153.08°E,;
rainforest, ca. 920 m, ex snl fragment
rotten wood (R. W. Taylor and R. Kohout ll
Mt. Chinghee, 12 km SE Rathdowney,
28.19°S, 152. 58°E, 720 m, rainforest, stick
brushing (Monteith, Yeates, and Thomp-|
son). New South Wales: Unumgar Forest
(Darlingtons); Woodenbong, rainforest
(Darlingtons); Tooloom Range, ca. 2,000 ft
(Darlingtons): Mt. Warning, Painforeem
800 to 3,500 ft, under and heaven rocks/
nest in wat rotten log but 2 inches below
ground (B. B. Lowery); Mt. Warning, 10
fa from Murwillumbah, RF, ca. 3,000 ft,
between rocks (B. B. Lowery); Whianj
Whian S. F., 28.39°S, 153.20°E, rainforest,
200 m, under stone, ace. no. 16993
16.vi.1976 (P. Ward); Tomewin, rainforest.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
1500 ft, under stone (B. B. Lowery); Blue
Knob Mt., Nightcap Ranges, rainforest,
2,800 to 3,000 ft (B. B. Lowery); Bilambil,
N of Tumbulgum, rainforest (B. B. Low-
ery); Hills above Tumbulgum, 800 ft, near
Murwillumbah, rainforest (B. B. Lowery);
Bonalbo, Sandy Crk., RF, 3000 ft, under
rock (E. G. Kearney).
Ecological Information. This species oc-
curs in rainforest, nesting in the soil, usu-
ally under or between rocks, often in a tan-
gle of small plant roots; it probably re-
stricts its foraging activity to the soil and
leaf litter (Taylor, 1965, 1968).
An additional 11 specimens, including
nine workers, a queen, and an ergatoid
queen, are examined here. They show the
following some differences from the above
“examined material”: (1) The ventral cen-
ter of the clypeus possesses a_toothlike
prominence in these 11 specimens, but a
short transverse carina in the previous
“Material Examined” section; (2) in the
nine workers, the juncture between the
pronotum and the mesonotum bears sev-
eral short longitudinal rugae, varying from
superficial to rather distinct (but is smooth
and shining in P wheeleri); (3) in the
queen, the petiole node in profile is
wedge-shaped, and the propodeal spines
(ca. 0.13) are shorter than those (ca. 0.20—
0.24) in P. wheeleri; and (4) in the ergatoid
queen, only one ocellus is present; PSL1
and PSL2 are not shorter than 0.08 and
0.13, respectively (in P. wheeleri, three
ocelli are distinct; PSL1 and PSL2 are ca.
0.18 and ca. 0.25, respectively). Further
collecting and studying will help deter-
mine whether these differences are signif-
icant or not.
These 11 specimens have the following
measurements: Worker: TL 3.50—4.76, HL
0.90-1.05, HW 0.94—1.12, CI 104—108, SL
Woo—l.01, SI 89=-93. EL 0.16—0.20, PW
0.60-0.68, AL 0.8S—0.96, PSL1 0.10—-0.14,
ES2 0.16-0.20, PPW 0.24-029 PPL
0.21—0.23, PPI 104-126 (n = 9). Queen:
Wi 3:84, Mi 1.22, HW 1.32) CI 108, SL
Ov, SL 8, Bi 0:25; PW 0:74, AL. 1.18,
PSL2 0.13, PPW 0.36, PPL 0.28, PPI 129
481
(a = IL), laigseicorel Giueeing INL, Sic4) Is0b,
Oo, Jay IOs) (Gib Ike}, Sib; Oyetss, SIL UL
EL 0.20, PW 0.64, AL 0.94, PSL1 0.08,
IE SIERA (OS). IRIAN (O75) IRIEL, (OL PARAL JIS)
(n = 1).
Collecting Data for the 11 Specimens
(ANIC). Australia: New South Wales,
Tuckers Knob, 21 km SW Coffs. Hbr.,
rainforest, 760 m, ANIC Berlesate No. 201
(N. I. Mitchell); New South Wales, Bellan-
gry, F'st, rainforest, ca. 3000 ft, iv.1958
(Darlingtons); New South Wales, Up. Al-
lyn R., near Eccleston, 32.08°S, 151.29°E,
rainforest, 400 m, acc. no. 316 (P. Ward):
New South Wales, Upr. Allyn Val., near
Eccleston, rainforest, ca. 2,000 ft, fallen
epiphyte masses, ANIC Berlesate 45, 11—
14.xii.1967 (Taylor and Brooks); New
South Wales, Comboyne plat., 2 to 2,800
ft, under rocks, x.1957 (Darlingtons).
Pristomyrmex wilsoni Taylor
Figures 193-194
Pristomyrmex wilsoni Taylor, 1968: 63. Holotype
worker, Australia: N. Queensland, Mt. Lewis, ca.
3,000 ft, near Julatten, rainforest, ground strays,
30—31.x.1966 (R. W. Taylor) [ANIC (Taylor, 1968)];
[one paratype worker (MCZC) examined].
Diagnosis (Worker). Pronotal spines and
propodeal spines exceptionally long (ca.
0.66—0.88).
Worker. TL 4.14-4.92, HL 0.96-1.11,
HW 0.96-1.12, Cl 96—104, SL 1.21—1.46,
SM 2a sos OM9=02 2 PVWE0:65—0N76:
AL Ada 3s05 PRPWr02S=0:32] PRE 030=
ORIEL 9 300K — 3)
Mandibles usually smooth and shining
but sometimes with one to two longitudi-
nal rugae. Masticatory margin of mandible
with three teeth arranged as an apical + a
preapical + a long diastema + a broad,
truncated basal tooth. Basal margin of
mandible lacking a distinctly curved lobe
or tooth. Clypeus usually with a median
longitudinal carina, but sometimes this
median carina is interrupted or indistinct.
Anterior clypeal margin with a median
denticle and two others on each side, but
sometimes two lateral denticles are fused
into one prominence. Ventral surface of
482
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 193-194. Pristomyrmex wilsoni Taylor. 193A: Worker head, full-face view; 193B: Showing a ruga on the ventral clypeus; -
194: Worker, lateral view.
clypeus with a short transverse carina. Palp
formula 2,3. Frontal carinae short, not be-
yond, or just reaching to the level of the
posterior margins of eyes; sometimes fron-
tal carinae absent Antennal scrobes ab-
sent. Frontal lobes absent; thus, the anten-
nal articulations are entirely exposed. An-
tennal scapes long, when laid on the dorsal
head, surpassing the occipital margin by
about one-third of their length. Eyes con-
taining 10 to 11 Ommnatichnk: in the longest
row. Eronile shape of alitrunk and pedicel
segments as in Figure 194. Pronotal spines
exceptionally long (ca. 0.66—0.88), curved
at their apices, diver ging toward the out-
sides in dorsal view. Propodeal spines ex-
ceptionally long (ca. 0.66—0.82), as illus-
trated in Figure 194; in dorsal view, they
are somewhat joined together at the base
but are divergent posteriorly, Metapleural
lobes small-triangular, each with an apex.
Petiole with a fairly long anterior pedun-
cle. Petiole node and postpetiole in profile
higher than long, in dorsal view each with
a somewhat aontenl apex. Postpetiole in
dorsal view usually longer than broad,
broadening from front to Thal Dorsum of
head usually smooth and shining, except
for a few foveolate punctures present on
the genae and sometimes bordering the
frontal carinae. Dorsum of alitrunk un-!
sculptured and highly polished. Petiole,
postpetiole, and gaster smooth and shine
ing. Dorsal surfaces of head and alitrunk
with numerous erect or suberect long.
hairs. A pair of similar long hairs bilaterally
distributed on the dorsum of petiole node
and on the postpetiole, respectively. First
gastral tergite lacking erect or suberect:
hairs. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with some erect
to suberect short hairs. Color reddish-
brown to blackish-brown.
Queen and Male. Unknown.
Comments and Discussion. Pristomyr- |
mex wilsoni occurs only in North Queens-’
land, Australia. It is a unique species in the:
genus that possesses both exceptionally:
long pronotal and propodeal spines and.
Thee can be immediately recognized. |
Pristomyrmex wilsoni may abe derived
from the common ancestor of the clade
consisting of P. curvulus and P. longispinus
of the Philippines. The workers of P. wil-)
soni are similar to those of P. curvulus and.
P. longispinus in possession of long pro-|
notal spines and long antennal scapes and
in the size and sculpture of body. Pristo-|
myrmex wilsoni differs from P. curvulus:
REVISION OF THE ANT GENUS PRISTOMYRMEX *® Wang
and P. longispinus in the workers as fol-
lows:
P. wilsoni
Propodeal spines exceptionally long, ca.
0.66 to 0.82; in dorsal view, they are
close to each other at the base
Maxillary palp with two segments
Masticatory margin of mandible with a
broad, truncated basal tooth
Petiole node and postpetiole, in dorsal
view each with a somewhat conical
apex
A pair of hairs present near the apex of
petiole node
P. curvulus and P. longispinus
Propodeal spines short to moderately
long, ca. 0.12 to 0.26; in dorsal view,
they are separated at the base
Maxillary palp with one segment
Masticatory margin of mandible with
two small basal teeth
Petiole node and postpetiole in dorsal
view each lacking a conical apex
Two or more pairs of hairs present on
the dorsal surface of petiole node
An alternative is that P. wilsoni might
evolved from the ancestor of the Austra-
lian P. wheeleri because of the following
characters being similar: (1) masticatory
margin of mandible with three teeth, (2)
maxillary palp with two segments, (3) body
smooth, and (4) ventral surface of clypeus
with a short transverse carina. However,
the workers of P. wilsoni obviously differ
from those of P. wheeleri as follows:
P. wilsoni
Pronotal and propodeal spines excep-
tionally long (ca. 0.66—0.88)
Antennal scapes relatively long (SL
1.21—1.46; SI 125-133)
Petiole node and postpetiole in dorsal
view each with a conical apex
Dorsum of alitrunk unsculptured
Labial palp with three segments
P. wheeleri
Pronotal and propodeal spines moder-
ately long (ca. 0.12—0.28)
483
Antennal scapes relatively short (SL
0.94—1.18; SI 8S—100)
Petiole node and postpetiole in dorsal
view lacking a conical apex
A transverse ridge present at the ap-
proximate position of metanotal
groove
Labial palp with two segments
Material Examined (ANIC, MCZC).
Australia: N. Queensland, Mt. Lewis,
1,000 m, RF (R. W. Taylor); Queensland,
Mt. Lewis, 960 m, 16.35°S, 145.17°E, rain-
forest, acc. no. 76.349 (R. W. Taylor and
T. A. Weir); NE Queensland, 2.5 km N
Mt. Lewis via Julatten, 1,040 m, RF, Py-
rethrum knockdown (D. K. Yeates and G.
I. Thompson); N. Queensland, 2 km SE
Mt. Spurgeon via Mt. Carbine, 1,100 m,
Pyrethrum tree logs (Montelth and
Thompson); NE Queensland, McDowall
Ra, 17 km N Daintree, 16.06°S, 145.20°E,
rainforest, 520 m, sieved litter, QM Ber-
lesate No. 684 (G. Montelth); NE Queens-
land, Table Mt., 10 km S of Cape Tribu-
lation, 16.09°S, 145.26°E, rainforest, 320
m, sieved litter, QM Berlesate No. 542 (G.
B. Montelth and D. Cook).
Ecological Information. Taylor (1968)
reports that this species occurs in rainfo-
rest; it has been collected at an elevation
of about 915 m in little berlesates and on
the surfaces of rocks and logs. “Collections
were made on two overcast days but not
on a third, which was brightly sunny.
Strays were not seen during several hours
night collecting (to about four hours after
sunset) at the same locality” (Taylor, 1968).
THE TRISPINOSUS GROUP
Worker. Medium-sized ants (HL 0.8S—
JED OFSP ILS, Wi, B24) satin
the following combination of characters:
(1) Masticatory margin of mandibles
with four teeth that have one of the fol-
lowing two arrangements:
a. the strongest apical + the second
strongest preapical + a relatively
short (first) diastema + a small den-
ticle + a relatively long (second) di-
484
Figures 195-196. Pristomyrmex bispinosus (Donisthorpe). 195:
Propodeal spines, dorsal view.
astema + a small basal denticle, as
in P. bispinosus and P. trispinosus (in
P. trispinosus, sometimes the first di-
astema is somewhat indistinct) or
b. the apical + the preapical + a rela-
tively long (first) diastema + a small
denticle + a relatively short (second)
diastema + a small basal denticle, as
in P. browni; sometimes the second
diastema is indistinct.
(2) Anterior margin of the median por-
tion of clypeus with at least five denticles.
Two ends of the anterior clypeal margin
each with a developed subtriangular tooth.
Lateral portions of clypeus reduced to a
margin; thus, the antennal fossae reach the
anterior clypeal margin.
(3) Palp formula 1,2.
(4) Frontal carinae absent.
(5) Frontal lobes almost completely ab-
sent.
(6) Antennal scrobes absent.
(7) Lamella that encircles the base of
antennal scape entire.
(8) Dorsum of alitrunk with a prome-
sonotal suture or impression.
(9) Alitrunk in profile with a convex
promesonotum and a deeply concave pro-
podeal dorsum.
(10) Pronotum with a pair of robust,
short to moderately long spines.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 |
Worker head, full-face view; 196A: Worker, lateral view; 196B: |
(11) Propodeal spines long, in dorsal !
view joined together at the base so that
they form a fork. |
(12) Petiole with a long anterior pedun-
cle that is longer than the length of the
node. |
(13) Foveolate punctures completely
absent, but regular striations are present
on the dorsal surfaces of the head and the
alitrunk in P. trispinosus and P. browni and
present on the genae and around the an-_
tennal sockets in P. bispinosus.
(14) First gastral tergite with numerous |
hairs.
This is a monophyletic group because it |
possesses many autapomorphic characters,
such as characters 3, 4, 8, 9, 11, and 13.
This group contains three native Mauritian —
species.
Pristomyrmex bispinosus (Donisthorpe)
Figures 195-196
Dodous bispinosus Donisthorpe, 1949: 272. Lecto-
type worker, Mauritius: Le Pouce Mt., 2.xi.1948 (R.
Mamet) (BMNH), here designated, [examined].
Pristomyrmex bispinosus (Donisthorpe) Brown, -
1971: 3.
Diagnosis (Worker). Dorsal surfaces of |
head and alitrunk mostly unsculptured;
mesonotum unarmed, at most weakly tu- —
berculate; alitrunk in profile with a convex |
REVISION OF THE ANT GENUS PRISTOMYRMEX ° Wang
promesonotum and a deeply concave pro-
podeal dorsum; propodeal spines in dorsal
view forming a divergent fork.
Worker. TL 4.36—4.82, HL 1.14—1.20,
IW 1.08—1.16, Cl 90=100, SL 1.32=—1.40,
Sl 18=130), EL 0:22=0'245 PW 0:70=0.75;
AL, 1.14—1.20; PPW 0:28-0:32, PPI 0.26—
ODS TARO syC a Lib)e
Mandibles smooth and shining. Masti-
catory margin of mandible with four teeth
arranged as the strongest apical + the sec-
ond strongest preapical + a relatively short
diastema + a small denticle + a relatively
long diastema + a small basal denticle.
Basal margin of mandible lacking a tooth-
like prominence. Anterior margin of the
median portion of clypeus with at least five
denticles, but sometimes two lateral den-
ticles are fused into a broad, truncated
lobe. Two ends of anterior clypeal margin
each with a strong, subtriangular tooth.
Ventral surface of clypeus usually with a
transverse ruga. Clypeus usually unsculp-
tured, but frontal area usually with three
to four short carinae that often extend to
the posterior clypeal margin. Palp formula
1,2. Frontal carinae absent. Antennal scro-
bes absent. Frontal lobes nearly complete-
ly absent so that the antennal articulations
are almost entirely exposed. Antennal
scapes long, when lying on the dorsal head
surpassing the occipital margin by about
one-fourth of their length. ‘Eyes usually
containing 11 to 12 ommatidia in the lon-
gest row. Occipital margin slightly concave.
Alitrunk in profile with a convex pro-me-
sonotum and a deeply concave propodeal
dorsum. Pronotum armed with a pair of
fairly long, acute spines. Propodeum with
a pair of long spines that, in dorsal view,
are joined together at the base but are di-
vergent along their length so that they
form a fork (Fig. 196B). Mesonotum un-
armed but sometimes with tubercles. Pro-
mesonotal suture or impression present.
Metapleural lobes dentiform. Petiole in
profile with a long anterior peduncle; an-
terodorsal angle of the node is higher than
the posterodorsal. Postpetiole in profile
usually with a curved anterior and dorsal
485
surface, in dorsal view slightly broader
than long. Dorsum of head mostly smooth
and shining but with some regular short
rugae present around the antennal fossae,
on genae and sometimes around the cen-
trical disc. Dorsum of alitrunk as well as
the sides of pronotum usually smooth and
shining, but the sides of the rest of alitrunk
usually with some regular short rugae. Pet-
iole unsculptured. Dorsum of postpetiole
smooth and shining. Gaster unsculptured.
All dorsal surfaces with numerous erect or
suberect hairs. A row of forward-project-
ing hairs present near the anterior clypeal
margin. Scapes and tibiae with some short
hairs. Color yellow-brown, but sometimes
reddish-brown.
Queen and Male. Unknown.
Comments. This species occurs in Maur-
itius. It can be easily separated from its
two relatives, P. trispinosus and P. browni,
in the workers, as follows: In P. bispinosus,
the dorsal surfaces of the head and the al-
itrunk are smooth and shining, except for
some short rugae around the atennal fos-
sae, on the genae, and sometimes around
the cenineal disc of the head: but in P
trispinosus and P. browni, the dorsal sur-
faces of the head and the alitrunk are en-
tirely sculptured with regular striations.
Material Examined (MCZC, ANIC,
MNHN). Mauritius: Le Pouce Mt., 700 to
SOO m, native forest (W. L. Brown); Le
Pouce, 700 m, 20°12’S, 57°31'E, ex closed
forest, on low vegetation (P. S. Ward); Le
Pouce Mt. (Ray Mamet).
Ecological Information. This species oc-
curs in a forest and has been collected on
the trees and on the main path (Brown,
1971).
Pristomyrmex browni sp. n.
Figures 197-198
Diagnosis (Worker). Dorsal surfaces of
head and alitrunk with dense regular stri-
ations; propodeal spines in dorsal view
joined together at base and subparallel
along their length; HW 0.82—0.90 and HL
0.88—1.01.
486
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 197-198. Pristomyrmex browni sp. n. 197: Worker head, full-face view; 198A: Worker, lateral view; 198B: Propodeal
spines, dorsal view.
Holotype Worker (MCZC). TL. 3.50, HL
0.92, HW 0.84, CI 91, SL 0.88, SI 105,
PW 0.46, AL 0.90. Paratypes, 11 workers
and one male (MCZC, ANIC, BMNH).
Worker. TL 3.42-3.78, HL 0.92-1.00,
HW 0.82-0.90, CI 87-94, SL 0.86—0.97,
SIR9S= Ee Os14= O19 RW 04420550;
AL 0.84—0.96, PPW 0.26—0.28, PPL 0.22—
Oy IRIE IZ ae ip = ILL).
Mandibles smooth and shining. Masti-
catory margin of mandible with four teeth
arranged as the strongest apical + the sec-
ond strongest preapical + a relatively long
(first) diastema + a small denticle + a rel-
atively short (second) diastema + a small
basal denticle: sometimes the second dia-
stema indistinct. Basal margin of mandible
almost straight, lacking a toothlike promi-
nence. Anterior margin of the median por-
tion of clypeus with at least five toothlike
prominences, but sometimes two lateral
denticles are fused into a broad lobe. Two
ends of the anterior clypeal margin each
with a strong subtriangular tooth. Ventral
surface of clypeus lacking toothlike prom-
inences. Clypeus usually with a few weak
longitudinal rugae. Palp formula 1,2.
Frontal carinae absent. Antennal scrobes
absent. Frontal lobes nearly completely
absent so that the antennal articulations
are almost entirely exposed. Antennal
scapes, when lying on the dorsal head,
slightly surpassing the occipital margin by
about one-eighth to one-seventh of their
length. Eyes moderately sized. Occipital
margin medially deeply emarginate. Ali-
trunk in profile with a convex pro-meson-
otum and a deeply concave propodeal dor-
sum. Pronotum with a pair of acute short
spines. Mesonotum lacking spines or teeth
but usually with three blunt small tuber- |
cles that are present on the posterior end |
and on the two sides, respectively. Propo--
deal spines in profile well developed, long,
bent at about a right angle near the base.
In dorsal view, propodeal spines somewhat |
laterally compressed; they are joined to-
gether at the base but subparallel along_
their length (Fig. 198B). Metapleural |
lobes small, triangular. Petiole node in |
profile nodiform with a long anterior pe- |
duncle; its anterodorsal angle is higher
than the posterodorsal. Postpetiole in pro- |
file with a single curved anterior and dor- |
sal surface, in dorsal view slightly broader |
than long and also broader than the petiole /
node. Dorsum of head entirely covered |
with regular coarse striations that consist
of longitudinal rugae and a few rugae |
around the antennal fossae. Sides and dor- |
sum of alitrunk sculptured with coarse cir-
cular striations evenly, but the center of
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
mesonotum with a few coarse longitudinal
rugae. Sides of petiole and postpetiole
usually with a few superficial rugae. Gaster
smooth and shining. Dorsal surfaces of
head, alitrunk, petiole, and postpetiole
with some erect or suberect hairs. First
gastral tergite with numerous recumbent
hairs. A row of forward-projecting hairs
present near the anterior clypeal margin.
Scapes and tibiae with numerous suberect
short hairs. Color reddish-brown.
Queen. Unknown.
Male. One paratype male, collected in
Mauritius by W. L. Brown, was originally
mounted together with a worker on the
same pin: TL 3.04, HL 0.58, HW 0.48, CI
So. OL) 0.20) SI 42, HWE 0:76, EL 0:38,
PW 0.73, AL 1.08, PPW 0.24, PPL 0.20,
RPT 120m = 1).
Head, including the eyes, broader than
long. Clypeus convex without a median
longitudinal carina. Anterior clypeal mar-
gin transverse. Frontal carinae absent.
Scapes only slightly longer than the first
funicular segments but distinctly shorter
than the rest of the funicular segments. On
the mesoscutum, notauli distinct, showing
a V shape; parapsidal furrows very super-
ficial. Scuto-scutellar sulcus with six nar-
row ridges. Propodeum weakly tubercu-
late, lacking teeth or spines. Metapleural
lobes triangular. Middle and hind tibiae
without any spurs. Petiole node in profile
with a fairly long anterior peduncle; ante-
rior face of the node, together with the
dorsal surface of the peduncle, forming a
declivity. Postpetiole in profile low and
rounded dorsally and in dorsal view broad-
er than long. Dorsum of head smooth and
shining, but frontal area with a median
longitudinal carina. Pronotum and mesos-
cutum rather smooth and shining, except
for those marked sutures, but mesoscutel-
lum with some longitudinal and reticulate
rugae. Petiole smooth and shining but with
a longitudinal carina present on each side
of the dorsal surface. Postpetiole and gas-
ter smooth and shining. All dorsal surfaces
with abundant erect or suberect hairs.
Scapes and tibiae with numerous erect or
487
suberect short hairs. Color reddish-brown:
wings slightly smoky.
Comments. This new species occurs in
Mauritius and Reunion Island. It is closely
related to the Mauritian P. bispinosus and
P. trispinosus. Characters separating P.
browni from P. bispinosus and P. trispino-
sus are provided under P. bispinosus and
P. trispinosus, respectively. The following
additional characters should also be men-
tioned: In the workers of P. browni, the
sculpture of the dorsal head consists most-
ly of longitudinal rugae, except for a few
rugae around the antennal fossae; the two
sides of the mesonotum each has a small
tubercle. In P. trispinosus, many striations
present on the dorsal head spread out
from the center to the two sides; the me-
sonotum possesses a pair of strong digitlike
prominences. In the type worker speci-
mens of P. browni, the first gastral tergite
possesses only recumbent hairs, and the
occipital margin of the head in full-face
view is deeply concave, which are different
may sz bispinosus and P. trispinosus.
One non-type specimen shows the fol-
lowing variations: (1) The first gastral ter-
gite possesses some erect or suberect
hairs, (2) the occipital margin of the head
in full-face view is feebly concave, (3) the
sides of the mesonotum lack any tubercles,
(4) the two small basal denticles on the
masticatory margin of the mandible are in-
distinct, (5) the dorsum of the petiole pe-
duncle in profile is obviously curved, and
(6) the rugae are very superficial on the
clypeus and on the sides of the petiole.
This specimen has the following measure-
ments: TL 3.24, HL 0.88, HW 0.84, CI 95,
SIO SOs SIN 9os HIE OMA ENV Ox4 05s AE:
OM ave
Holotype Worker. Mauritius: Le Pouce
Mt., 700 to SOO m, native forest, 1.iv.1969
(W. L. Brown).
Paratypes. Mauritius: Le Pouce Mt.,
700 to 800 m, native forest, 4.iii. 1977 (W.
L. Brown).
Additional Non-Type Worker Examined.
Indian Ocean, Reunion I., Mare Longue,
488
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 199-200. Pristomyrmex trispinosus (Donisthorpe). 199:
Propodeal spines, dorsal view.
near St. Philippe, primitive forest, 580 m,
15.i.1975 (D. Schauenberg) (MCZC).
Ecological Information. This species oc-
curs in native forest.
Pristomyrmex trispinosus (Donisthorpe)
Figures 199-200
Dodous trispinosus Donisthorpe, 1946: 145. Syntype
workers and males, Mauritius: Cocotte Mt.,
27.xii.1941 and 6.ii.1943 (R. Mamet) (BMNH,
LACM, MCZC) [examined].
Pristomyrmex trispinosus (Donisthorpe) Brown,
1971: 3.
Diagnosis ( Worker). Pronotum, meson-
otum, and propodeum each with a pair of
strong prominences; dorsal surfaces of
head and alitrunk with dense regular stri-
ations; propodeal spines in dorsal view di-
vergent; HW and HL > 1.10.
Worker. TL 4.50-4.68, HL 1.16—1.22,
EIW. eS ess Cl Sl=97 Sie 133 18445
SI17/=122 EL 020201245, PW 0:66=0570:
AL 1.16—1.26, PPW 0.27-0.30, PPL 0.24—
0:26) RPI 108=115 Ga — 6).
Mandibles smooth and shining. Masti-
catory margin of mandible with four teeth
arranged as the strongest apical + the sec-
ond strongest preapical + a short diastema
(sometimes, this diastema is not distinct)
+ a small denticle + a long diastema + a
small basal denticle. Basal margin of man-
dible lacking a toothlike prominence. An-
terior margin of the median portion of
Worker head, full-face view; 200A: Worker, lateral view; 200B: |
clypeus with at least five denticles, but
sometimes two lateral denticles are fused
into a broad lobe. Two ends of the anterior -
clypeal margin each with a developed
prominence. Ventral surface of clypeus
lacking toothlike prominences. Clypeus—
usually with a few longitudinal rugae. Palp—
formula 1,2. Frontal carinae absent. An-
tennal scrobes absent. Frontal lobes ab- |
sent so that the antennal articulations are
completely exposed. Antennal scapes long, -
when lying on the dorsal head surpassing |
the occipital margin by one-fourth to one-_
third of their length. Eyes usually contain-—
fi
ing 11 to 12 ommatidia in the longest row. |
Occipital margin in full-face view slightly |
concave. Alitrunk in profile with a convex
pro-mesonotum and a deeply concave pro-
podeal dorsum. Pronotum armed with a
pair of moderately long acute spines. Me-
sonotum with a pair of thick, blunt, digit-
like short prominences. Propodeum with a_
pair of developed long spines that, in dor-
sal view, are joined together at the base
but divergent along their length so that
they form a fork (Fig. 200B). Metapleural |
lobes dentiform. Petiole node in profile |
nodiform with a long anterior peduncle. i
Postpetiole in profile rounded anterodor-
sally but usually with a distinct postero-
dorsal angle; in dorsal view, postpetiole -
slightly broader than long. Entire dorsum —
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
of head with regular coarse striations:
many striations spread out from the center
to the two sides and to the occipital mar-
gin; some are around the antennal fossae,
the rest are some longitudinal rugae pre-
sent on the sides of the head. Sides and
the dorsum of alitrunk with numerous
evenly distributed, circular coarse stria-
tions. Center of mesonotum with a few
coarse short rugae. Petiole and postpetiole
rather smooth and shining, but sometimes
their sides with a few superficial short ru-
gae. Gaster unsculptured. All dorsal sur-
faces with numerous erect or suberect
hairs. Scapes and tibiae with numerous
erect or suberect short hairs. A row of for-
ward-projecting hairs present near the an-
terior clypeal margin. Color yellow-brown,
but sometimes reddish-brown.
Queen. Unknown.
Male. One syntype male (BMNH), to-
gether with a number of syntype workers,
constitutes a series (see Donisthorpe,
1946): TL 4.46, HL 0.72, HW 0.70, CI 97,
SL 0.20, SI 29, EL 0.22, PW 0.90, AL 1.42
im = 1].
Head, including the eyes, broader than
long. Clypeus somewhat transverse, con-
vex in middle. On the mesoscutum, notauli
indistinct. Scuto-scutellar sulcus wide, sep-
arated into small cells by narrow ridges.
Propodeum weakly tuberculate, lacking
teeth or spines. Metapleural lobes sub-
triangular. Petiole node in profile low with
a fairly long anterior peduncle. Postpetiole
in profile rounded dorsally. Dorsum of
head smooth and shining. Mesoscutum
smooth and shining, but mesoscutellum
with some rugae and a few foveolate punc-
tures. Petiole node rather smooth. Post-
petiole and gaster unsculptured, smooth,
and shining. All dorsal surfaces with abun-
dant erect or suberect long hairs. Scapes
and tibiae with numerous erect or sub-
erect short hairs. Color reddish-brown:
wings somewhat dusky.
Comments. Pristomyrmex trispinosus is
known only from Mauritius. It differs from
P. bispinosus in the workers in having reg-
ular coarse striations on the entire dorsal
489
surfaces of the head and the alitrunk and
a pair of strong, digitlike prominences on
the mesonotum. The workers of P. trispi-
nosus and P. bispinosus are separable from
those of P. browni by the following char-
acters:
P. trispinosus and P. bispinosus
Propodeal spines in dorsal view diver-
gent, not laterally compressed; in pro-
file rather straight
A relatively short diastema present be-
tween the preapical and the third
tooth on the masticatory margin of
the mandible
Larger species with HW > 1.08, HL >
ae Sie oe NN OO Sale =
4.36
P. browni
Propodeal spines in dorsal view subpar-
allel, somewhat laterally compressed;
in profile, bent at about a right-angle
near the base
A relatively long diastema present be-
tween the preapical and the third
tooth on the masticatory margin of
the mandible
Smaller species with HW 0.82-0.90, HL
0.88—1.01, SL 0.80—0.97, PW 0.40—
ODO 324= 3.05
Material Examined (MCZC). Mauritius:
Cocotte Mt. (R. Mamet).
Ecological Information. A nest of P. tri-
spinosus was found under a flat stone, and
about 30 to 40 workers, two males, larvae,
and pupae were collected; this species,
when disturbed, simulates death (Donis-
thorpe, 1946).
THE LEVIGATUS GROUP
Worker. This group shows the following
combination of characters:
(1) Usually small-sized species: HW:
0.60—0.96 in 11 species, 0.98—1.26 in one
species (P. lucidus). HL: 0.60-0.90 in 11
species, 0.92—1.16 in P. lucidus. TL: 2.20-
3.49 in 11 species, 3.71—4.84 in P. lucidus.
(2) Masticatory margin of mandible
with four teeth arranged as the strongest
apical + the second strongest preapical +
490
the smallest third + an acute basal tooth
that is larger than the third tooth but
smaller than the apical and preapical teeth;
masticatory margin lacking a distinct dia-
stema.
(3) A tooth that is short and broad or
prominent, present about midway on the
basal margin of mandible.
(4) Lateral portions of clypeus in front
of antennal insertions reduced to a narrow
margin.
(5) Lamella that encircles the base of
antennal scape entire.
(6) Anterior clypeal margin with at
most three denticles (i.e., a median den-
ticle and one on each side), but sometimes
the median tooth is indistinct so that only
two denticles are seen on the margin.
(7) Ventral surface of clypeus smooth or
bearing a weak transverse ruga but lacking
toothlike prominences.
(8) Palp formula 1,3 in 10 species ex-
amined.
(9) Frontal carinae extending to the lev-
el of the posterior margins of eyes.
(10) Well-developed scrobes absent.
(11) Dorsum of alitrunk in profile more
or less arched, in dorsal view without any
sutures.
(12) Pronotum usually unarmed but
with a pair of small teeth in one species
(P. minusculus).
(13) Petiole node in profile high, higher
than the length of the node, usually with
a distinct anterior face, but in P. inermis,
the anterior face of the petiole node is in-
separable from the dorsal surface of the
peduncle.
(14) Dorsal surfaces of head and ali-
trunk smooth or with scattered foveolate
punctures or with foveolate-reticulate
sculpture.
This is a monophyletic group, contain-
ing 12 species. They are endemic in the
Oriental region, except for one species (P.
minusculus) entering in North Queens-
land, Australia. Further, most of the spe-
cies of this group occur in Papua New
Guinea and some nearby islands.
Obviously, this group, together with the
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
profundus group, constitutes a monophy-
letic lineage because the workers and
queens of the two groups possess a syna- |
pomorphy, that is, the masticatory margin
of the mandible with four teeth (the stron-—
gest apical + the second strongest preap-
ical + the smallest third + an acute basal |
tooth) but lacking a diastema. The workers
of the group are easily distinguished from
those of the profundus group by lacking
well-developed antennal scrobes and pos-
sessing (1) two to three teeth on the an-
terior clypeal margin, (2) a more or less |
arched dorsum of the alitrunk, and (3) a
tooth present about midway on the basal
margin of the mandible but not adjacent
to the basal tooth of the masticatory mar-
gin.
The dentition of the masticatory margin
of the mandible in the workers and queens
of the levigatus group is a critical character
separating the levigatus group from other
five (i.e., cribrarius, punctatus, quadri-—
dens, and umbripennis)
groups.
In the workers and queens of the levi-
gatus group, foveolate punctures show
trispinosus,
continuous variation on the dorsum of the |
head between the frontal carinae. I treat
this case as follows: (1) P. levigatus almost |
completely lacks distinct foveolate punc-
tures on the dorsum of the head between
the frontal carinae; (2) P. simplex assem-
bles those populations with some scattered
foveolate punctures, but spaces between
foveolae are usually smooth; and (3) P. cog-
vii shows foveolate-reticulate sculpture;
some populations, only with foveolate-re-
ticulate sculpture behind the eyes, are
considered intermediate forms and also
grouped into P. coggii. In fact, this similar
continuous variation also occurs in the oth-
er two (i.e., guadridens and umbripennis)
groups.
An ergatoid queen caste has been found
in P. mandibularis of the group. This caste —
is also present in the punctatus and quad-
ridens groups. Pronotal armaments, widely
occurring in the cribrarius, quadridens,
and trispinosus groups, appear in the -
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
49]
Figures 201-202. Pristomyrmex acerosus sp. n. 201: Worker head, full-face view; 202: Worker, lateral view.
workers of one species (P. minusculus) of
the levigatus group. These facts indicate
that ergatoid queens and pronotal arma-
ments have arisen several times in Pristo-
myrmex, respectively.
Pristomyrmex acerosus sp. n.
Figures 201-202
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
subpetiole with a pinlike long process.
Holotype Worker (BMHH). TL. 2.80,
HL 0.76, HW 0.75, CI 99, SL 0.66, SI 88,
EL 0.12, PW 0.48, AL 0.76, PPW 0.22,
REE OM. PPI W29)
Mandibles smooth and shining. A
broad-based short tooth present about
midway on the basal margin of the man-
dible. Clypeus depressed, unsculptured,
its anterior margin with three denticles: a
median tooth and one on each side. Fron-
tal carinae just extending to the level of
the posterior margins of eyes. Antennal
scrobes absent. Frontal lobes weakly ex-
panded basally. Eyes moderately sized.
Occipital margin feebly concave in full-
face view. Dorsum of alitrunk in profile
convex. Pronotum unarmed. Propodeum
with a pair of subtriangular short spines.
Metapleural lobes rounded. Petiole node
in profile high, with the anterodorsal angle
higher than the posterodorsal, its anterior
face subparallel to the posterior one, an-
terior peduncle of the node about as long
as the node. Subpetiole with a semitrans-
lucent pinlike long process. Postpetiole
with a rounded dorsum. In dorsal view,
petiole node subrounded; postpetiole
broader than long, somewhat transrectan-
gular. Dorsal surfaces of head and alitrunk
smooth and shining, except for some small,
shallow hair pits. Petiole, postpetiole, and
gaster unsculptured, smooth, and shining.
Dorsal surface of head with numerous
erect to suberect hairs. Dorsum of alitrunk
with sparse hairs. Two pairs of hairs pre-
sent on the dorsum of petiole node. A few
on the dorsum of postpetiole and on the
first gastral tergite, respectively. Three
pairs of forward-projecting hairs present
near the anterior clypeal margin. Scapes
and tibiae with some suberect short hairs.
Color lightly yellow-brown.
Queen and Male. Unknown.
Comments and Discussion. This is the
only species in the genus that has so far
been found to have a semitranslucent, pin-
like, long process on the ventral surface of
the petiole. However, further collecting is
required. If this long process is present in
other specimens, this holotype represents
a good species; otherwise, this holotype
492
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 203-204. Pristomyrmex boltoni sp. n. 203: Worker head, full-face view; 204: Worker, lateral view.
would be an aberrant specimen, and P. ac-
erosus would become a junior synonym of
P. levigatus.
Holotype Worker. New Hebrides: Ma-
lekoula I. (N), Vao Isl, 0 to 20 m, 7.ix.1979
(W.C. Gagne, G. M. Nishida, and G. A.
Samuelson).
Ecological Information. Unknown.
Pristomyrmex boltoni sp. n.
Figures 203-204
Diagnosis ( Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
eyes very small, with two to three omma-
tidia in the longest row; dorsal surfaces of
head between frontal carinae and alitrunk
smooth and shining.
Holotype Worker (LAMN). TL 2.48, HL
0.66, HW 0.66, CI 100, SL 0.52, SI 79,
EL 0.08, PW 0.42, AL 0.62, PPW 0.18,
PPL 0.16, PPI 113. Paratypes, two workers
(LAMN, MCZC).
Worker. TL 2.40, 2.40; HL 0.65, 0.66;
HW 0.66, 0.66; CI 100, 102; SL 0.55, 0.55:
SI 83, 83; EL 0.08, 0.08; PW 0.44, 0.44;
AL 0.60, 0.61; PPW 0.18, 0.18; PPL 0.16,
(IS IPI WIE TLS Go =D).
Mandibles smooth and shining but with
few longitudinal rugae in the paratypes. A |
broad-based short tooth present about —
midway on the basal margin of the man- |
dible. Frontal area concave with a median
carina. Clypeus flat, unsculptured, smooth, —
and shining, its anterior margin with two —
lateral teeth, but in the two paratypes, an
additional weak median tooth present. |
Frontal carinae distinct, extending to the
level of the posterior margins of eyes.
Scrobal impressions shallow, present lat- |
eral to the frontal carinae in full-face view. —
Frontal lobes weak so that the antennal ar- —
ticulations are almost entirely exposed. An- |
tennal scapes, when lying on the head, |
close to the occipital margin. Eyes very
small, with two to three ommatidia in the
longest row. Pronotum unarmed. Propo- |
deum with a pair of triangular teeth. Me-_
tapleural lobes rounded. Petiole node in —
profile high with the anterodorsal angle
higher than the posterodorsal, its anterior —
peduncle about as long as the node. Post- |
petiole in profile higher than long with a
rounded dorsum. In dorsal view, petiole —
node subrounded, about as broad as long;
postpetiole broader than long. Dorsum of —
head between the frontal carinae smooth |
and shining but with some foveolate punc- —
tures present around the eyes. Dorsum of |
REVISION OF THE ANT GENUS PRISTOMYRMEX ° Wang
alitrunk smooth and shining. Petiole and
postpetiole smooth and shining. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with numerous erect or suberect
hairs. Two pairs of hairs present on the
dorsum of petiole node, three pairs on the
dorsum of postpetiole, and a few on the
base of the first gastral tergite. A few pairs
of forward-projecting hairs present near
the anterior clypeal margin. Scapes and
tibiae with some erect or suberect short
hairs. Color reddish-brown.
Queen and Male. Unknown.
Comments. Pristomyrmex boltoni is a
sibling species of P. longus and P. coggii.
The three species are all from New Guin-
ea. Their workers possess very small eyes.
Pristomyrmex boltoni differs from P. lon-
gus and P. coggii as follows: The dorsum
of the petiole node in dorsal view is slight-
ly broader than long or about as broad as
long in the workers of P. boltoni, but long-
oval and distinctly longer than broad in P.
longus. The dorsal surfaces of the head be-
tween the frontal carinae and the alitrunk
are smooth in the workers of P. boltoni but
are covered with numerous foveolate
punctures in P. coggii.
The workers of P. boltoni are also similar
in appearance to those of P. levigatus, but
they can be separated by the following
characters:
P. boltoni
Eyes smaller, with the maximum diam-
eter 0.08, containing two to three om-
matidia in the longest row
P. levigatus
Eyes larger, with the maximum diame-
ter 0.12 to 0.16 (rarely 0.10), contain-
ing five to seven ommatidia in the
longest row
Holotype Worker. New Guinea: Gulf
Prov., Ivimka Camp, Lakekamu Basin,
7.73°S, 146.76°E, 120 m, #96-235, lowland
wet forest, ex sifted leaf litter, 28.x.1996
(R. R. Snelling).
Paratypes. Two workers, New Guinea:
Gulf Prov., Ivimka Camp, Lakekamu Ba-
sin, 7.7°S, 146.8°E, 140 m elevation, #96-
493
280, lowland wet forest, ex sifted leaf litter,
6.xi.1996 (R. R. Snelling).
A non-type specimen (ANIC), collected
in New Guinea (Brown R., lowland RF,
under log) by B. B. Lowery, has the fol-
lowing measurements: HW 0.66, HL 0.66,
SL 0.56, EL 0.08, PW 0.44, AL 0.60.
Ecological Information. This species oc-
curs in lowland forest and has been col-
lected in litter samples.
Pristomyrmex coggii Emery
Figures 205-208
Pristomyrmex coggii Emery, 1897: 584. Lectotype
worker, New Guinea: Montes Hansemanni et Ber-
linhafen (Bird) (MCSN), here designated, [exam-
ined].
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
eyes with two to four ommatidia in the
longest row; dorsal surface of head, except
for the scrobal areas, with foveolate-retic-
ulate sculpture; dorsum of petiole node in
dorsal view about as long as broad or
slightly broader than long; first gastral ter-
gite with only a few hairs.
Worker. TL 2.20-2.86, HL 0.60-0.74,
HW 0.60-0.76, CI 98-104, SL 0.48—0.64,
SI 74-86, EL 0.05—0.09, PW 0.40-0.48,
AL 0.60-0.77, PPW 0.16—0.20, PPL 0.12—
Of SRP aS Sr33e(n— 2.0).
Mandibles usually smooth and shining
but sometimes with a few superficial lon-
gitudinal rugae. A broad and short or
strongly prominent tooth present about
midway on the basal margin of mandible.
Clypeus flat, its anterior margin sometimes
with three denticles: a median denticle
and one on each side, but sometimes the
median denticle absent or weak so that
only two teeth are present there. Frontal
area concave, with a median carina that
usually extends to the clypeus. Ventral
center of clypeus lacking any rugae or
prominences. Palp formula 1,3. Frontal ca-
rinae distinct, extending to the level of the
posterior margins of eyes. Scrobal impres-
sions smooth and shallow, present lateral
494
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 205-208. Pristomyrmex coggii Emery. 205: Worker head, full-face view; 206: Worker, lateral view; 207: Queen head,
full-face view; 208: Queen, lateral view.
to the frontal carinae. Frontal lobes weakly
expanded so that the antennal articulations
are almost completely exposed. Eyes
small, usually with two to three, rarely
four, ommatidia in the longest row. Occip-
ital margin straight or feebly concave in
full-face view. Pronotum unarmed. Pro-
podeum with a pair of triangular short
spines. Metapleural lobes rounded. Petiole
node in profile high, with the anterodorsal
angle higher than the posterodorsal, its an-
terior peduncle about as long as the node.
In dorsal view, dorsum of petiole node
subrounded, about as long as broad, or
transoval, slightly broader than long. Sub-
petiole with a narrow, long, semitranslu-
cent lamella. Postpetiole in profile round-
ed dorsally, in dorsal view somewhat trans- |
verse-rectangular and broader than long. —
Dorsum of head, except for the scrobal ar- _
eas, with dense foveolate punctures that |
form foveolate-reticulate sculpture; some- _
times the punctures are almost aligned so —
that it seems that the several longitudinal —
rugae appear between the frontal carinae.
Dorsum of alitrunk with sparse foveolate _
punctures. Petiole, postpetiole, and gaster —
unsculptured, smooth, and shining. Dorsal _
surfaces of head and alitrunk with numer-
ous erect to suberect hairs. Dorsal surfaces
of petiole node and _ postpetiole usually —
with two pairs of hairs, respectively. A few |
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 495
hairs present on the base of the first gastral
tergite. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with numerous
erect to suberect short hairs. Color red-
dish-brown.
Queen. TL 2.94-3.16, HL 0.72-0.78,
HW 0.72-0.82, CI 100—105, SL 0.56—0.66,
Si Sone en Ola —OslivmueNViOlaa—OKO4-
ANlEnOM 80.96, BPW 021-024" PPL.OllW—
OalSs PPL 1222133 (n= 3).
Generally similar to worker, except for
normal caste differences. In addition, fo-
veolate punctures shallow on the meson-
otum, propodeal armaments slightly short-
er than those in conspecific worker.
Male. Unknown.
Comments and Discussion. Pristomyr-
mex coggii is closely related to P. boltoni
and P. longs. The three species occur in
New Guinea. Characters separating P. cog-
gii from P. boltoni are provided unde: fie
latter name. Pristomyrmex coggii differs
from P. longus because the deisuin of the
petiole Roden in dorsal view is about as long
as broad or broader than long in the wor ia
ers of P. coggii but longer than broad in P.
longus.
Pristomyrmex coggii differs from P. obe-
sus of Solomon Islands as follows: The
workers of P. coggii possess only a few
hairs on the first gastral tergite and have
smaller eyes containing two to three, rarely
four, ommatidia in the longest row. But in
the workers of PB. obesus, the entire first
gastral tergite is evenly covered with erect
or suberect hairs, and the eyes contain five
to seven (rarely four) ommatidia in the
longest row. Pristomyrmex coggii differ
from P. simplex of New Guinea and the
Philippines because the dorsum of the
head between the frontal carinae bears fo-
veolate-reticulate sculpture in the workers
of P. coggii but only scattered foveolate
punctures in P. simplex; in addition, the
eyes usually contain five ommatidia in the
longest row in the workers of P. simplex
Pit - usually two to three in P. coggii.
It must be pointed out that then material
I have examined may resolve into two spe-
cies with further study. Additional collect-
ing will help clarify the situation.
Material Examined (MCZC, ANIC,
LAMN, BMNH, NHMYV). Papua New
Guinea: Gulf Prov., Ivimka Camp, Lake-
kamu Basin, lowland wet forest, #96-291
@273"S. TA6./67E) 120) msex sifted leafilit=
ter), #96-350 (7.7°S, 146.8°E, 400 m ele-
vation, sifted leaf litter and debris from
rotten log) (R. R. Snelling); W. Highlands,
Baiyer R., 4,000 ft (S. Peck); Rinona (R.
Vane-Wright); New Guinea (Bird); Nad-
zab, dry evergreen forest (E. O. Wilson);
Huon Pen., lower Busu R., lowland rain-
forest (E. O. Wilson); Wau, Kilolo Creek
(J. Balogh); Lae, rainforest, leaf mold (T.
E. Woodward); Lae, Busu R., lowland
rainforest, in rotten wood on ground (B
B. Lowery); NW Lae, “Timber Track” ca.
16 km, rainforest, ca. 220 m, berlesate (R.
W. Taylor); near Popondetta, <50 m (R.
W. Taylor); 8 km S of Kokoda, 800 m, rain-
forest (R. W. Taylor).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter samples and in rotten wood on the
ground.
Note: I have examined additional 25
specimens, including 19 workers and 6
queens. In these workers, the eyes are
slightly larger than those in P. coggii; they
contain four to five ommatidia in the lon-
gest row. Further collecting and studying
are needed.
These 25 specimens have the following
measurements: Worker: TL 2.76—3.00, HL
0.73—0.80, HW 0.73-0.82, CI 100-106, SL
0.58-0.64, SI 74-82, EL 0.09-0.10, PW
0.48—0.52, AL 0.68—0.82 (n = 19). Queen:
2) 92235345) Elle OM2—O0FS 0S EIN OM/2=
0.82, CI 100-103, SL 0.56—0.62, SI 73-82,
I CLIO IEW O50. G2. ALL O.SO=
0.92 (n = 6).
Collecting Data for the 25 Specimens
(WI GAG: ANIC. BMNH). New Guinea:
Morobe Dist., Bulolo (B. B. Lowery); Bul-
olo, rainforest, 3,500 ft (B. B. Lowery);
NE New Guinea, 6 km NE of Wau, Bulolo
R. valley, rainforest, 1,100 m, berlesate,
leaf mold (R. W. Taylor); Wau, Bishop Mu-
496
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
(2
KM
Figures 209-212. Pristomyrmex inermis sp. n. 209: Worker head, full-face view; 210: Worker, lateral view; 211: Queen head,
full-face view; 212: Queen, lateral view.
seum Station, 1,200 m, rainforest, rotten
log (R. W. Taylor); Wau, 4,000 ft, forest
litter (S. Peck); N. Wau, on Bulolo Rd.,
650 m (S. Peck); Wau, Kunai Creek, rain-
forest, ca. 1,400 m, berlesate (R. W. Tay-
lor); Tapini, 1,000 to 1,200 m, rainforest,
rotten log (R. W. Taylor).
Pristomyrmex inermis sp. n.
Figures 209-212
Diagnosis (Worker). Propodeum lacking
a pair of teeth or spines.
Holotype Worker (LAMN). TL 2.86, HL
0.72, HW 0.70, CI 97, SL 0.64, SI 91, EL
0.14, PW 0.50, AL 0.72. Paratypes, 17
workers and three queens (ANIC, LACM,
MCZC, BMNH).
Worker. TL 2.65-—3.36, HL 0.70-0.84,
HW 0.69-0.84, CI 96-103, SL 0.62—0.76,
SI 83-92, EL 0.12-0.16, PW 0.46-0.58,
Al? 0.63-0:80; PPW 0:23-027 Reb Ove
O21 PRI 12821425(n— ls)
Mandibles smooth and shining but
sometimes with a few superficial small
punctures. Masticatory margin of mandi-
ble with four teeth arranged as the stron-
gest apical + the second strongest preap-
ical + the smallest third + a basal tooth;
a distinct diastema lacking. A broad-based
triangular short tooth present about mid-
way on the basal margin of mandible.
Frontal area concave. Clypeus flat,
smooth, and shining, its anterior margin
with three denticles: a weak median tooth
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
and one on each side, but sometimes the
median tooth indistinct or lacking so that
only two teeth are present there. Ventral
surface of clypeus lacking any distinct ru-
gae or teeth. Palp for vile 1,3. Frontal ca-
rinae just extending to the ewe of the pos-
terior margins of fhe eyes. Antennal scro-
bes absent. Frontal lobes very weak. Eyes
moderately sized. Occipital margin feebly
convex in full face view. Dorsum of ali-
trunk in profile arched. Pronotum un-
armed. Propodeum lacking a pair of teeth
or spines but usually with a pair of blunt
small tubercles. Metapleural lobes small,
usually bluntly rounded, but sometimes
toothlike. Petiole node in profile wedge-
shaped, usually with a blunt triangular
apex; sometimes the apex somewhat blunt-
ly rounded. Subpetiole with a narrow rim.
Postpetiole in profile slightly higher than
the petiole node, with a convex dorsum.
In dorsal view, petiole node and postpeti-
ole broader than long. Cephalic dorsum
between the frontal carinae highly pol-
ished but usually with a few foveolate
punctures bordering frontal carinae. A few
foveolate punctures present on the genae
and many on the ventral surface of the
head. Dorsum of alitrunk, petiole, and
postpetiole smooth and shining. Gaster
unsculptured. Dorsal surfaces of head, al-
itrunk, petiole, and postpetiole with sparse
erect to suberect hairs. A few hairs present
on the first gastral tergite. A few pairs of
forward-projecting hairs present near the
anterior clypeal margin. Scapes and tibiae
with some suberect hairs. Color varying
from light yellow-brown to blackish-
brown, gaster usually darker than alitrunk;
sometimes alitrunk lighter than head and
gaster so that the specimens show a bicol-
ored form.
Queen. TL 3.23-3.52, HL 0.80-0.81,
HW 0.76—0.85, CI 95-106, SL 0.66-—0.74,
SI 87-92, EL 0.18—0.20, PW 0.64—0.72,
AL 0.86—0.94, PPW 0.26—0.30, PPL 0.18—
O24 PPI 125-144 (m= 3):
General shape as in Figures 211-212;
except for normal caste differences, other
characters similar to worker.
497
Male. Unknown.
Comments. This species can be easily
recognized because it lacks a pair of pro-
podeal teeth or spines in the workers,
which is unique within Pristomyrmex. In
addition, in the levigatus group, P. inermis
has a characteristic shape of the petiole
node in the workers and queens: wedge-
like in profile view with an apex. This char-
acter has originated independently at least
twice in the genus because it is also seen
in P. fossulatus and P. punctatus of the
punctatus group.
Holotype Worker. Indonesia: Irian Jaya,
PT. Freeport Concession, Siewa Camp.,
03.04°S 136.38°E, 200 ft, lowland second-
ary rainforest, #98-71 (stray foragers in leaf
litter), 12.iv.1998 (R. R. Snelling).
Paratypes. One queen with same data as
holotype; one queen with same data as ho-
lotype but date 7.iv.1998, and number
#98-48 (stray on log); two workers with
same data as holotype but date 7.iv.1998
and number #98-44 (ex sifted leaf litter);
three workers, Papua New Guinea: Bulolo,
2.300 ft, rainforest, 19.xii. 1967 (B. B. Low-
ery); three workers, Papua New Guinea:
Bulolo, 2,300 ft, rainforest, 27.xii.1967 (B.
By Lowery); three workers, Papua New
Guinea: Bulolo, 2,600 ft, rainforest,
25.xii.1970 (B. B. Lowery); one worker,
Papua New Guinea: Bulolo, 2,800 ft, rain-
forest, Ji197l, (Be By Lowery); :thnee
workers, Papua New Guinea: Bulolo,
3,000 ft, rainforest, 5.i1.1971 (B. B. Low-
ery); one queen, New Guinea: Bulolo
(Morobe Dist), 8.i.1971 (B. B. Lowery);
two workers, New Guinea: near Vanimo,
rainforest, ca. 50 m, ex rotting log, 10—
11.vii.1972 (R.W.T.).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter samples and on logs.
Pristomyrmex largus sp. n.
Figures 213-214
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four ea, of which the third
foot counting from the apex, smallest:
495
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6 :
|
Figures 213-214. Pristomyrmex largus sp. n. 213: Worker head, full-face view; 214: Worker, lateral view.
HW and HL = 0.90; petiole node in pro-
file with the anterodorsal angle higher
than the posterodorsal.
Holotype Worker (ANIC). TL 3.38, HL
0.90, HW 0.96, CI 107, SL 0.78, SI 81,
EL 0.14, PW 0.61, AL 0.92, PPW 0.24,
PPL 0.20, PPI 120. Paratypes, 34 workers
and one queen (MCZC, R. Clouse’s per-
sonal collection).
Worker. TL. 3.40, HL 0.90, HW 0.90, CI
100, SL 0.81, SI 90, EL 0.14, PW 0.60,
AL 0.90, PPW 0.25, PPL 0.20, PPI 125 (n
= jl).
Mandibles smooth and shining. A
broad-based triangular short tooth present
about midway on the basal margin of the
mandible. Clypeus flat, unsculptured,
smooth, and shining; its anterior margin
with three denticles: a weak median tooth
and one prominent lateral tooth on each
side, sometimes the median tooth indis-
tinct. Ventral surface of clypeus unsculp-
tured and smooth. Palp formula 1,3. Fron-
tal carinae just extending to the level of
the posterior margins of eyes. Antennal
scrobes absent. Frontal lobes weakly ex-
panded. Eyes moderately sized. Occipital
margin straight or feebly concave in full-
face view. Pronotum unarmed. Propo-
deum with a pair of long spines that are
about 1.5 times the distance between their
bases. Metapleural lobes subtriangular.
Dorsum of alitrunk in dorsal view rather
flat. Petiole node in profile high, with the
anterodorsal angle higher than the poster-
odorsal, its anterior face sometimes sub-
parallel to the posterior one; anterior pe-
duncle of the node about as long as the
node, and subpetiole with a narrow semi-
translucent rim. In dorsal view, petiole
node transoval. Postpetiole in profile high-
er than long, rounded dorsally, in dorsal
view broader than long and somewhat
transrectangular. Both dorsal and ventral
surfaces of head smooth and shining but
with few small, shallow punctures present
on the genae. Dorsum of alitrunk smooth
and shining. Petiole and _ postpetiole
smooth and shining, except for a lateral
longitudinal carina on each side that sep-
arates the tergite from the sternite. Gaster
unsculptured. Dorsal surfaces of head and
alitrunk with some erect to suberect short
hairs. Petiole node and postpetiole each
with two pairs of hairs as illustrated in Fig-
ure 214. A few hairs present near the base
of the first gastral tergite. A few pairs of
forward-projecting hairs present near the
anterior clypeal margin. Scapes and tibiae
with abundant erect to suberect short
hairs. Color yellow-brown to reddish-
brown.
Queen. TL 3.94, HL 0.94, HW 0.95, CI
101, SL 0.80, SI 84, EL 0.22, PW 0.74,
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
AIG, TOI TA PWWEO BO). IPP O28) Ie ISO) a
= iDy
Generally similar to worker, except for
normal caste differences; in addition, pro-
podeum with a pair of short spines, shorter
than those in the conspecific worker, first
gaster tergite with numerous hairs.
Male. Gulonowns
Comments. Pristomyrmex largus is the
second largest species in the levigatus
group after P. lucidus. Pristomyrmex lar-
gus must have evolved from the ancestor
of P. levigatus. Pristomyrmex largus can be
separated from P. levigatus by ie follow-
ing characters in the vores:
P. largus
Larger species with HW = 0.90, and
HL = 0.90
Propodeal spines relatively long, longer
than the distance between en bases
Ventral surface of head smooth, with
only a few small hair pits
P. levigatus
Smaller species with HW < 0.80, and
HL < 0.80
Propodeal armaments relatively short,
shorter than or about as long as the
distance between their bases
Ventral surface of head with numerous
foveolate punctures
Holotype Worker. Ponape I., Mt. Tolen-
kiup; vi-ix.50 (P. A. Adams).
Paratypes. One worker, Micronesia:
Pohnpei I., Hilltop campsite near Mt.
Nanalaud, 400 m, on an old ivory nut,
3.v.1995 (Ron Clouse); 33 workers and one
queen, Pohnpei I., Nahnal aud cave,
around camp (#155—-158, under dead
leaves, on ground; #161, under rotting
leaves; #167, under dead leaves. gl iie= 180.
[eaves, ander rotten), 24.i1ii.2000 (Ron
Clouse).
Ecological Information. All par atypes of
this species have been collected in high-
elevation rainforest, under the rain- eoalked
leaf litter, and inside a rotten ivory nut on
the forest floor (R. Clouse, personal com-
munication).
Note: I do not illustrate the queen of
499
this species because this caste, together
with 33 workers, collected by Mr. Ronald
Clouse, reached me after my manuscript
was completed.
Pristomyrmex levigatus Emery
Figures 215-218, 277, 281
Pristomyrmex levigatus Emery, 1897: 583. Lectotype
worker, New Guinea: near Friedrich-Wilhelmshaf-
en et Berlinhafen (Biré) (MCSN), here designated,
[examined].
Pristomyrmex mendanai Mann, 1919: 341. Two syn-
type workers, Solomon Is.: Santa Cruz Is., Graciosa
Bay, 19.v.-24.xi.1916 (W. M. Mann) (MCZC,
USNM) [examined]. Syn. n.
Note: The lectotype of P. levigatus
shows some fine differences from the two
syntypes of P. mendanai. In the former, the
metapleural lobes are subtriangular; the
dorsum of the alitrunk is rather flat, with
two sides somewhat margined; the pro-
podeum is armed with a pair of short
spines that are not subtriangular but some-
what incurved at their apices. In the latter,
the metapleural lobes are rounded; the
dorsum of the alitrunk is convex, and its
sides are not distinctly margined; the pro-
podeal armaments are subtriangular, not
distinctly incurved at their apices. How-
ever, the presence of some intermediate
forms suggests that they belong to the
same species for the present.
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
HW 0.62-0. 78, HL 0.64—0.78; eyes with
five to eight ommatidia in the longest row;
pronotum unarmed; propodeum with a
pair of teeth or short spines; petiole with
a lateral longitudinal carina on each side;
subpetiole lacking a pinlike long process;
dorsal surfaces of head and _ alitrunk
smooth and shining.
Worker. TL 2 36-2 .96, HL 0.64—0.78,
HW 0.62-0.78, CI 96-106, SL 0.50—0.65,
SI 78—89, EL 0.12—0.16 (very rarely 0.10),
PW 0.41-—0.52, AL 0.54—0.75, PPW 0.20—
O22 RR ON4=OMSaeRe i223 1435 ae
30).
500
217
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
‘ons
N
Figures 215-218. Pristomyrmex levigatus Emery. 215: Worker head, full-face view; 216: Worker, lateral view; 217: Queen
head, full-face view; 218: Queen, lateral view.
Mandibles usually smooth and shining
but with a few fine longitudinal rugae in
some specimens. A broad-based triangular
short tooth present about midway on the
basal margin of the mandible. Frontal area
concave, usually unsculptured, but some-
times with a weak median carina. Clypeus
flat, unsculptured, smooth, and shining; its
anterior margin with a median denticle
and two lateral teeth, but sometimes the
median tooth is weak or absent. Ventral
surface of clypeus lacking any rugae or
toothlike prominences. Palp formula 1,3.
Frontal carinae distinct, extending to the
level of the posterior margins of eyes.
Scrobal impressions shallow, present lat-
eral to the frontal carinae. Frontal lobes
weak so that the antennal articulations are
almost entirely exposed. Antennal scapes,
when lying on the head, close to the oc-
cipital margin. Eyes moderately sized, with
five to eight ommatidia in the longest row.
Dorsum of alitrunk in dorsal view flat in
the lectotype but convex in some speci-
mens. Pronotum unarmed or at most with
blunt tubercles, lacking teeth or spines.
Propodeum with a pair of triangular teeth
or short spines that are slender, slightly in-
curved at their apices in the lectotype. Me-
tapleural lobes subtriangular or rounded.
Petiole node in profile nodiform with the
anterodorsal angle higher than the poster-
odorsal; its anterior peduncle about as long
as the node. Subpetiole with a narrow long
flange. In dorsal view, the dorsum of pet-
iole node transoval and broader than long,
Postpetiole in profile higher than long,
with a rounded dorsum, in dorsal view
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
somewhat transverse-rectangular and
broader than long. Dorsum of head
smooth and shining but sometimes with a
few foveolate punctures bordering the
frontal carinae; sometimes a few foveolate
punctures present on the genae and
around the eyes. Dorsum of alitrunk un-
sculptured, smooth, and shining. Petiole
and postpetiole smooth and shining, each
with a longitudinal carina on each side that
separates the tergite from the sternite.
Gaster unsculptured. Dorsum of head
with numerous erect to suberect hairs.
Dorsum of alitrunk with some erect or
suberect hairs. Dorsal surfaces of petiole
node and postpetiole usually with one to
two pairs and one to three pairs of hairs,
respectively. A few hairs present at the
base of the first gastral tergite. A few pairs
of forward-projecting hairs present near
the anterior clypeal margin. Scapes and
tibiae with erect to suberect short hairs.
Color golden-brown or reddish-brown.
Queen. TL 2.92-3.34, HL 0.74—0.75,
HW 0.76—0.81, CI 101—108, SL 0.60—0.65,
SI 79-83, EL 0.18—0.19, PW 0.54—0.66,
AL 0.80—0.92, PPW 0.23-0.26, PPL 0.18—
0.20) PPI'128=133 @ = 5).
Generally similar to worker, except for
normal caste differences. Propodeal ar-
maments tuberculate or denticulate,
slightly shorter than those in the conspe-
cific worker.
Male (Figs. 277, 281). Two male speci-
mens, together with 13 workers and two
queens, collected in New Guinea (Nadzab,
#1083) by E. O. Wilson, constitute a se-
ries; each of the two males was originally
mounted together with a worker or with a
worker and a queen on the same pin: TL
2.48, 2.54; HL 0.50, 0.51; HW 0.53, 0.53;
1 105;,° 106: SL 0:16, 0.18: SI 30, 34:
HWE 0.63, 0.64; EL 0.23, 0.25; PW 0.60,
0.63; AL 0.82, 0.84: PPW 0.18, 0.19; PPL
0.15, 0.16; PPI 119, 120 (n = 2).
Head, including the eyes, broader than
long. Clypeus lacking a median longitudi-
nal carina; anterior clypeal margin rather
straight. Frontal carinae reaching the level
of the posterior margins of antennal inser-
501
tions. Mesoscutum lacking distinct notauli
and parapsidal furrows. Scuto-scutellar
sulcus with nine narrow longitudinal ridg-
es. Propodeum weakly tuberculate, lacking
teeth or spines. Metapleural lobes with a
blunt or somewhat rounded apex. Middle
and hind tibiae without any spurs. Petiole
node in profile low, nodiform, having an
anterior face and a long anterior peduncle.
Postpetiole in profile low and rounded
dorsally, in dorsal view subrectangular and
broader than long. Dorsum of head
smooth and shining, but frontal area with
a median longitudinal carina. Dorsal ali-
trunk generally smooth and shining, except
for those marked sutures. Dorsal surfaces
of petiole and postpetiole smooth and
shining. Gaster unsculptured. All dorsal
surfaces with abundant rather long hairs.
Legs and scapes with numerous erect or
suberect short hairs. Color blackish-
brown. Wings infuscated.
Comments and Discussion. Pristomyr-
mex levigatus occurs in New Guinea,
Nama Is., Solomon Is., New Georgia, and
New Britain Is. It appears to be a basal
species within the levigatus group. Many
species, such as P. acerosus, P. boltoni, P.
inermis, P. largus, P. lucidus, P. mandibu-
laris, P. minusculus, P. obesus, and P. sim-
plex, may have evolved from a P. levigatus—
like ancestor. The workers of these species
are separable from those of P. levigatus as
follows: P. levigatus differs from both P.
simplex and P. obesus by lacking foveolate
punctures on the dorsal surfaces of the al-
itrunk and the head between the frontal
carinae. In P. levigatus, the eyes are larger,
usually containing five to eight ommatidia
in the longest row but two to three in P.
boltoni. Pristomyrmex levigatus is smaller
(HW < 0.80, HL < 0.80) than P. largus
and P. lucidus (HW > 0.90, HL > 0. 90).
Pristomyrmex minusculus bears a pair of
pronotal teeth that are not seen in P. lev-
igatus. A pair of propodeal teeth or short
spines are present in P. levigatus, but ab-
sent in P. inermis. Pristomyrmex levigatus
possesses a longitudinal ruga on each side
of the petiole node that is not seen in P.
502
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 219-220. Pristomyrmex longus sp. n. 219: Worker head, full-face view; 220A: Showing the dorsum of the petiole node
of the worker, in dorsal view, is long-oval and longer than broad; 220B: Worker, lateral view.
mandibularis. Finally, the subpetiole of P.
levigatus does not have a pinlike long pro-
cess that is distinct in P. acerosus.
Material Examined (ANIC, BMNH,
LAMN, MCZC). New Guinea: Nadzab,
#1083, dry evergreen forest (E. O. Wil-
son); Gogol Val. ca. 24 km W. Madang, ca.
50 m, rainforest, rotten wood, ex small
fragment (R. W. Taylor); Bulolo, rainforest,
2,300 ft (B. B. Lowery); Yawasora near
Wewak, ca. 50 m, rainforest, berlesate (R.
W. Taylor); Gulf Prov., Ivimka Camp, Lak-
ekamu Basin, 7.73°S, 146.76°E, 110 m,
#96-345 (R. R. Snelling); Port Moresby,
Brown River (J. Baloph); N.D. Papua,
Sangara (G. Baker); (P. M. Room). Nama
Is. near Truk (R. W. L. Potts). Solomon Is.:
New Georgia (E. S. Brown). New Britain
Is. (L. Weatherill).
Ecological Information. This species oc-
curs in rainforest and has been collected
in a litter sample.
Pristomyrmex longus sp. n.
Figures 219-220
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
eyes small, with only two to three omma-
tidia in the longest row; dorsum of petiole
node in dorsal view long-oval and longer
than broad; first gastral tergite with nu-
merous hairs.
Holotype Worker (MCZC). TL 2.51, HL
0.67, HW 0.66, CI 99, SL 0.55, SI 83, EL
0.07, PW 0.44, AL 0.66. Paratypes, 14
workers (MCZC, ANIC, BMNH).
Worker. TL 2.22-2.68, HL 0.63—0.70;
HW 0.62-0.70, CI 95—100, SL 0.52—0.56,
SI 80-87, EL 0.05—0.10, PW 0.40-0.46
AL 0.60—0.70, PPW 0.17—0.18, PPL 0.16—
O16, PP 041 35G sla)»
Mandibles usually smooth and shining.
A broad-based short tooth present about
midway on the basal margin of the man-
dible. Clypeus flat, its anterior margin with
a median tooth and two lateral teeth;
sometimes the median tooth is weak.
Frontal area concave, with a median carina
that usually extends to the clypeus. Ventral
surface of clypeus smooth, lacking any ru-
gae or prominences. Palp formula 1,3.
Frontal carinae distinct, extending to the
level of the posterior margins of eyes.
Scrobal impressions shallow, smooth, pre-
sent lateral to the frontal carinae. Frontal
lobes weakly expanded. Antennal scapes,
when lying on the head, close to the oc-
cipital margin. Eyes very small, consisting
of 4 to 10 ommatidia, with only two to
three ommatidia in the longest row. Oc-
cipital margin straight or feebly concave in
full-face view. Profile of alitrunk and ped-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
icel segments as in Figure
tum unarmed. Propodeum with a pair of
triangular teeth. Metapleural lobes round-
ed. Betiole node in profile high with the
anterodorsal angle higher than the poster-
odorsal, its anterior fice subparallel to the
posterior one; anterior peduncle of the
node about as long as or slightly shorter
than the node; subpetiole with a narrow
semitranslucent lamella. In dorsal view,
dorsum of petiole node long-oval and dis-
tinctly longer than broad. ‘Postpetiole in
rofile rounded dorsally, in dorsal view
slightly broader than long or about as long
as broad. Dorsum of oad between the
frontal carinae mostly smooth but usually
with some sparse, small, and shallow punc-
tures. Some foveolate punctures present
on the genae and around the occipital cor-
ners of Mcadk Dorsum of alitrunk, petiole,
postpetiole, and gaster usually unsculp-
tured, smooth, anal shining. Dorsal surfac-
es of head, alitrunk, and gaster with nu-
merous erect to suberect haus. ‘Two pairs
of hairs usually present on the dorsal sur-
faces of petiole node and postpetiole, re-
spectively, as illustrated in Figure 220B. A
few pairs of forward-projecting hairs pres-
ent near the anterior clypeal margin.
Scapes and tibiae with numerous erect to
suberect short hairs. Color yellow-brown
to reddish-brown.
Queen and Male. Unknown.
Comments. The workers of P. longus are
quickly recognizable by the following: In
the levigatus group, only three species, P.
Polioni.: P. coggii, and P. longus, possess
small eyes. Only two species, P. longus and
P. obesus, have numerous hairs evenly dis-
tributed on the entire first gastral tergite.
But in only one species (P. onan) is ahve
dorsum of the petiole node in dorsal view
long-oval and longer than broad; in the
other members of the levigatus group, the
dorsum of petiole node is subrounded or
transverse-oval (i.e., about as long as broad
or broader than long).
Holotype Worker. New Guinea: Huon
Pen., Lower Busu R., lowl. rainfor.,
0.V.1955, #957 (E. O. Wilson).
220B. Prono-
503
Paratypes. Eight workers with same
data as holotype; one worker, New Guinea:
Huon Pen., Lower Busu R., lowland rain-
forest, 6.v.1955, #978 (E. O. Wilson); five
workers, New Guinea: 13 km NW Lae,
Bubia, lowland rainforest, 26.iii.1955, #688
(E. O. Wilson).
Ecological Information. This species has
been collected in lowland rainforest.
Pristomyrmex lucidus Emery
Figures 221-224
Pristomyrmex lucidus Emery, 1897: 584. Holotype
worker, New Guinea: Berlinhafen (Bird) (MCSN)
[examined].
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
HW 0.98-1. 26. HL 0.92-1.16; postpetiole
in dorsal view longer than broad or as long
as broad, in profile with an arched anterior
face and a steeply sloping posterior face;
petiole node in profile with a single evenly
rounded blunt apex.
Worker. TL 3:71-4.84, HE 0:92=1.16,;
HW 0.98-1.26, CI 104—111, SL 0.90—1.16,
SI 89-98, EL 0.16—0.20, PW 0.62-0.79,
AL 0.90—1.28, PPW 0.24—0.28, PPL 0.26-—
OSS RPLS4=100NGr — o>):
Mandible smooth and shining but
sometimes with a few longitudinal rugae
superficial or distinct. A broade based short
tooth present about midway on the basal
margin of the mandible. Clypeus with a
short median longitudinal carina that usu-
ally does not reach to the anterior clypeal
margin; sometimes this carina indistinct;
soniceimnes a few additional short rugae
present. Anterior clypeal margin wach a
median tooth and two lateral oe some-
times the median tooth is smaller than the
others. Ventral surface of clypeus lacking
any rugae or teeth. Palp formula 1,3. Fron-
tal carinae extending to the level of the
posterior margins of eyes. Antennal scro-
bes absent. F Frontal lobes : slightly expanded
basally. Eyes containing 7 to 10 ommatidia
in the longest row. reais of alitrunk and
pedicel segments as in Figure 222. Pron-
504
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 221-224. Pristomyrmex lucidus Emery. 221: Worker head, full-face view; 222: Worker, lateral view; 223: Queen head,
full-face view; 224: Queen, lateral view.
otum at most with a pair of blunt tuber-
cles, lacking teeth or spines. Propodeum
with a pair of armaments, varying from
broadly based minute teeth to moderately
long acute spines. Metapleural lobes tri-
angular or each with a_blunt-rounded
apex. Petiole node in profile high, with a
single evenly blunt-rounded apex and a
long anterior peduncle. Postpetiole in pro-
file high (slightly higher than petiole), with
an arched anterior face and a steeply slop-
ing posterior face, its apex pointing pos-
terior-upwardly. In dorsal view, postpetiole
broadening from front to back, mostly lon-
ger than broad, rarely about as long as
broad. Cephalic dorsum between the fron-
tal carinae highly polished but usually with
a few foveolate punctures bordering the
frontal carinae. Sometimes a few foveolate
punctures present on the genae and many
on the ventral surface of the head. Dor-
sum of alitrunk, petiole, and postpetiole
unsculptured and highly polished. Gaster
smooth and shining. Dorsal surfaces of
head and alitrunk with sparse erect to sub-
erect moderately long hairs. A pair of hairs
present on the dorsum of petiole node and
one to two pairs on the dorsum of post-
petiole. First gastral tergite with a few
hairs. A few pairs of forward-projecting
hairs present near the anterior clypeal
margin. Scapes and tibiae with numerous
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
suberect to erect short hairs. Body uni-
form yellow-brown or blackish-brown or
bicolored (i.e., alitrunk and pedicel seg-
ments lighter than head and gaster).
Oueen WWE oma a2. Sle A:
RIV 22 1 27 EL lOss Ile SE 106.413:
SI 87, 89; EL 0.26, 0.28; PW 0.96, 1.00;
Ee) 1240.42: -PPW 0:31) 0:36 PPE 0:32,
0.38; PPI 95, 97 (n = 2).
Generally similar to worker, except for
normal caste differences. In addition, pro-
podeal armaments usually shorter than
those in the conspecific worker; first gas-
tral tergite sometimes with numerous
erect or suberect hairs.
Male. Unknown.
Comments. The workers of P. lucidus
can be easily recognized by the following
characters: (1) The postpetiole, in profile
view, shows an arched anterior face and a
steeply sloping posterior face, with the
apex pointing posterior-upwardly. This
shape is unique in the levigatus group.
Furthermore, the postpetiole in dorsal
view is longer than or as long as broad. In
the other 11 species of the group, the post-
petiole, in dorsal view, is distinctly broader
than long. (2) The petiole node in profile
view bears a single evenly blunt-rounded
apex that is not seen in the other 11 spe-
cies of the levigatus group. (3) P. lucidus
has the largest head width (HW) in the
levigatus group: In the 55 specimens mea-
sured, HW is 1.00 to 1.26 but 0.98 in only
one individual. In the other species of the
levigatus group, HW is less than 0.90, ex-
cept in one species, P. largus, in which,
HW falls into the range 0.90 to 0.96.
Material Examined (ANIC, MCZC,
LAMN, USNM, CASC, BMNH#). Papua
New Guinea: Tapini, 1,000 to 1,200 m,
rainforest, acc. 2249 (rtw. wood fragment),
acc. 2252 and 2262 (nest in soil under log)
(R. W. Taylor); near Vanimo, rainforest, ca.
50 m (ex rotting log) (R. W. Taylor); Ya-
wasora near Wewak, <50 m, rainforest, ex
rotting wood piece (R. W. Taylor); 6 km
NE of Wau, Bulolo R. Valley, rainforest
1,100 m, under bark of rotten log (R. W.
Taylor); 8 km S of Kokoda, rainforest, 800
505
m, vial 4-36 (ground strays) and vial 37-
191 (ex rotting wood piece) (R. W. Taylor);
Wau, Bishop Museum Station, 1,200 m, ex
soil under rotten branch (R. W. Taylor); 9
mi on Lae, side of Mumeng, 3,500 ft, rain-
forest (B. B. Lowery); Bulolo, 4,000 ft,
rainforest (B. B. Lowery); Bulolo Gorge,
2,800 ft, rainforest (B. B. Lowery); along
Kokoda Rd., 400 to 1,000 ft, rainforest (B.
B. Lowery); 2 mi N Kokoda, ca. 1,000 ft,
rainforest (B. B. Lowery); Wau Gorge,
3,000 ft, rainforest (B. B. Lowery); Wau,
4,000 ft, rainforest (B. B. Lowery); ca. 12
km SE Vanimo, 150 m Virgin hill rainfo-
rest (W. L. Brown); Wau N on Bulolo Rd.
B-278, 650 m (S. Peck); Bewani Rd., near
Vanimo km 2 quarry, 40 m, lowland rain-
forest (W. L. Brown); Maffin Bay (E. S.
Ross). Indonesia: Irian Jaya, PT. Freeport
Concession, Wapoga Camp., 03.14°S,
136.57°E, 3,450 ft, #98-230 (Montane pri-
mary rainforest, ex rotten stick in litter) (R.
R. Snelling).
Ecological Information. This species oc-
curs in rainforest and has been collected
in soil under a log, under the bark of a
rotten log, in litter samples, and on the
ground.
Pristomyrmex mandibularis Mann
Figures 225-228
Pristomyrmex mandibularis Mann, 1921: 444. Syn-
type workers, Fiji Is.: Viti Levu, Nadarivatu, 1915—
1916 (W. M. Mann) (AMNH, LACM, MCZC,
USNH) [examined].
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest; a
strongly prominent tooth present about
midway on the basal margin of mandible;
petiole lacking a lateral longitudinal carina
on each side; dorsum of head between
frontal carinae usually smooth and shining.
Worker. TL 2.68—-3.49, HL 0.74—0.84,
HW 0.76-0.88, CI 95-107, SL 0.61—0.74,
SI 80-93, EL 0.11—0.14, PW 0.48—0.60,
AL 0.68—0.90, PPW 0.20-0.25, PPL 0.16—
O19] RPMs — se)!
Mandibles usually smooth and shining.
506 Bulletin Museum
of Comparative Zoology, Vol. 157, No. 6
Figures 225-228. Pristomyrmex mandibularis Mann. 225: Worker head, full-face view; 226: Worker, lateral view; 227: Ergatoid
queen head, full-face view; 228A: Ergatoid queen, lateral view; 228B: Dorsum of the ergatoid queen alitrunk, dorsal view.
A prominent tooth present about midway
on the basal margin of mandible. Clypeus
flat and unsculptured. Anterior clypeal
margin with a median tooth and two lat-
eral teeth; the median tooth, in size, sim-
ilar to or smaller than the others. Ventral
surface of clypeus usually with a weak
transverse ruga. Palp formula 1,3. Frontal
carinae extending to the level of the pos-
terior margins of the eyes. Antennal scro-
bes absent. Frontal lobes slightly expand-
ed. Eyes moderately sized. Occipital mar-
gin straight or feebly concave in full-face
view. Pronotum unarmed. Propodeum
with a pair of short to moderately long
spines. Metapleural lobes triangular. Peti-
ole node in profile high, higher than long,
with the anterodorsal angle being an apex
and the dorsum posteriorly rounding into
the posterior surface, its anterior peduncle
slightly longer than or about as long as the
node. Postpetiole in profile high, about
two times as high as long, rounded dor-
sally. In dorsal view, postpetiole transrec-
tangular. Dorsum of head between the
frontal carinae usually smooth and shining
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
but sometimes with a few small and shal-
low punctures. Sometimes a few foveolate
punctures bordering the frontal carinae,
present on the genae and around the eyes.
Dorsum of alitrunk smooth and shining.
Petiole and postpetiole smooth and shin-
ing, each lacking a lateral longitudinal ca-
rina on each side. Gaster unsculptured.
Dorsal surface of head with numerous
erect to suberect long hairs. Some of sim-
ilar hairs present on the dorsum of ali-
trunk, a pair on the petiole, one to two
pairs on the postpetiole, and a few on the
first gastral tergite. A few pairs of forward-
projecting hairs present near the anterior
clypeal margin. Scapes and tibiae with nu-
merous erect to suberect hairs. Color red-
dish-brown to blackish-brown.
Ergatoid Queen. TL 3.32, 3.40; HL
0.84, 0.84; HW 0.82, 0.82; CI 98, 98; SL
0.66, 0.73; SI 80, 89; EL 0.14, 0.15; PW
0.54, 0.56; AL 0.78, 0.82; PPW 0.23, 0.24;
PIPIE, OS). OfVile IPP TS a Ge, =)
Generally similar to worker, color and
pilosity as in the worker, but the head with
one ocellus, alitrunk in dorsal view with a
pro-mesonotal suture, mesonotum more
convex, and propodeal armaments shorter
than in the conspecific worker.
Queen and Male. Unknown.
Comments. Pristomyrmex mandibularis
is endemic in Fiji and is the only Pristo-
myrmex species so far occurring there. It
possesses an ergatoid queen caste, which
is not seen in the other species of the lev-
igatus group; furthermore, its ergatoid
queen has only one ocellus.
Pristomyrmex mandibularis is closely
related to P. levigatus and P. largus, also
from the Oriental region. The differences
between the workers of P. mandibularis
and those of P. levigatus and P. largus are
as follows:
P. mandibularis
Petiole and postpetiole each lacking a
lateral longitudinal carina on each
side
Basal margin of mandible with a strong-
ly prominent tooth
507
Anterior clypeal margin usually with a
distinct median tooth
P. levigatus and P. largus
Petiole and postpetiole each with a lon-
gitudinal carina on each side that sep-
arates the tergite from the sternite
Basal margin of mandible with a broad-
based short tooth
A median tooth, on the anterior clypeal
margin, usually lacking or very weak
Material Examined (ANIC, MCZC,
USNM, BMNH, MHNG). Fiji: Nausori
Highlands, #424 (rotting stick in litter) (W.
L. and D. E. Brown); Viti, Nadarivatu,
rainforest, acc. 83 (berlesate, leaf mold)
and acc. 66.51 (forest floor, colony in small
crevice, rotting branch fragment) (R. W.
Taylor); Nadarivatu (W. M. Mann); Lase-
ma (W. M. Mann); Viti, Levu, Nadarivatu
Reserve, 17.34'S, 177.57'E, Rainforest 800
m, Q. M. Berlesate No. 775, sieved litter
(G. Monteith); Kadavu, 2 km SE Vunisea,
19.04’'S, 178.10'E, rainforest 20 m, Q. M.
Berlesate No. 770, sieved litter (G. Mon-
teith).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter berlesates; it nests beneath stones
in small colonies (Mann, 1921).
Pristomyrmex minusculus sp. n.
Figures 229-232
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and pos-
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
pronotum with a pair of teeth.
Holotype Worker (MCZC). TL 3.02, HL
0.76, HW 0.80, CI 105, SL 0.66, SI 83,
EL 0.14, PW 0.52, AL 0.78. Paratypes, 16
workers and one queen (MCZC, ANIC,
BMNH, NACA).
Worker. TL 2.52-3.02, HL 0.66—0.78,
HW 0.66-0.80, CI 99-106, SL 0.54—0.66,
SI 77-84, EL 0.10-0.14, PW 0.45-0.52,
AL 0.59-0.74, PPW 0.20-0.24, PPL 0.14—
0.19, PPI 126-143 (n = 16).
Mandibles usually smooth and shining
but sometimes with a few small shallow
508
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 229-232. Pristomyrmex minusculus sp. n. 229: Worker head, full-face view; 230: Worker, lateral view; 231: Queen
head, full-face view; 232: Queen, lateral view.
punctures. A broad-based short tooth pre-
sent about midway on the basal margin of
the mandible. Clypeus unsculptured. An-
terior clypeal margin with a median tooth
and two lateral teeth, but the median tooth
often smaller than the others, sometimes
the median tooth indistinct. Ventral sur-
face of clypeus with a weak transverse
ruga. Palp formula 1,3. Frontal carinae ex-
tending to the level of the posterior mar-
gins of the eyes. Scrobal areas shallow,
short, present lateral to the frontal carinae
in full-face view. Frontal lobes weakly ex-
panded. Eyes moderately sized. Occipital
margin feebly concave. Alitrunk in dorsal
view more or less flat. Pronotum with a
pair of acute small teeth; in some small
specimens, this pair of teeth are very weak
but visible, and in dorsal view they become
a pair of sharp points on the two sides of
the pronotum. Propodeum armed with a
pair of spines, varying in length and shape,
straight to slightly upcurved along their
length. Metapleural lobes subtriangular,
but rarely with a rounded apex. Petiole
node in profile high with the anterodorsal
angle higher than the posterodorsal, its an-
terior surface usually subparallel to the
posterior one, its anterior peduncle about
as long as the node. Subpetiole with a nar-
row rim. Postpetiole in profile high,
rounded dorsally. Petiole node and post-
petiole in dorsal view broader than long.
Dorsum of head between the frontal ca-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
rinae smooth and shining but sometimes
with a few foveolate punctures bordering
the frontal carinae, present on the genae
and around the eyes. Dorsum of alitrunk
smooth and shining. Petiole and postpeti-
ole each with a lateral longitudinal ruga on
each side that separates the tergite from
the sternite. Gaster unsculptured. Dorsal
surfaces of head and alitrunk with sparse
erect to suberect hairs. Dorsal surfaces of
petiole node and postpetiole each usually
with two pairs of hairs and first gastral ter-
gite with a few hairs, as in Figure 230. A
few pairs of forward-projecting hairs pre-
sent near the anterior clypeal margin.
Scapes and tibiae with some erect to sub-
erect short hairs. Color yellow-brown to
reddish-brown; appendages lighter.
Queen. TL 3.08, HL 0.72, HW 0.74, CI
103, SL 0.60, SI 81, EL 0.17, PW 0.60,
AL 0.82, PPW 0.24, PPL 0.18, PPI 133 (n
= 1)
Generally similar to worker, except for
normal caste differences. In addition, pro-
notal armaments absent; propodeum with
a pair of teeth or short spines that are
shorter than those in the conspecific work-
er.
Male. Unknown.
Comments. This species has a wide dis-
tribution in the Oriental region. It is also
dispersed to North Queensland, Australia.
Pristomyrmex minusculus must be derived
from a P. levigatus—like ancestor. It is in-
distinguishable, in the queen, from P. lev-
igatus at present. The workers of P. min-
usculus have almost same appearance as
those of P. levigatus, except for a derived
character—the pronotum with a pair of
small teeth. In the levigatus group, this
critical character is possessed only by the
workers of P. minusculus, thus making
them easily recognizable.
Holotype Worker. Palau Is: Peleliu L.,
east coast, 26.1.1948 (H. S. Dybas).
Paratypes. Nine workers and one queen
with same data as holotype; one worker,
Wallis Is.: NukuTapu I., 28.iii.1965 to
l.iv.1965 (G. Hunt). Two workers, Indo-
nesia: Seram, above Haruru, near Masohi,
509
50 to 150 m,
one worker, Indonesia: Irian Jaya, 12 km
S of Sorong, forest fragment, 1.v.1981 (W.
L. Brown); two workers, Tonga Is.: Fale-
hau, Niuatoputapu, moss + lichen, from
coconut tree trunks, 1.ix.1971 (W. and G.
Rogers); two workers, YapGroup, vii—
viii.50. (R. J. Goss).
The following additional (non-type)
specimens have a pair of very weak pro-
notal prominences. They have the follow-
ing measurements: Worker: HW _ 0.60—
0.72, HL 0.62—0.72, CI 97-103, SL 0.52—
0.60, SI 77-87, EL 0.10-0.12, PW 0.42-—
0.48, AL 0.57—0.73, PPW 0.19—-0.21, PPL
0.14—0:116, PPI 125-150 (n = 12). Oueen:
IED Ow Ol Moe Ini; One, OLwes (Cll IC)
LOOSE GO!625 0622 SIeS2aro2 ken OokS:
0.18; PW 0.62, 0.62; AL 0.82, 0.88; PPW
ORAS 0:24 SRP OM ORI eA Ss Aa
(a= 2).
Collecting Data for These Non-Type
Specimens (ANIC. USNM, BMNH). Papua
New Guinea: Kiunga (J. Balogh); Bisian-
umu near Sogeri, rainforest, 500 m (E. O.
Wilson); Maffin Bay (E. S. Ross). Micro-
nesia: Pohnpei, Agric. and Trade School,
in leaf litter (in Ylang- Ylang grove) and in
rotting coconut tree (Ron Clouse). Austra-
lia: N. Queensland, Cape York, Lockerbie
(G. B. Monteith); N.Q., Iron Ra. rainfo-
rest, berlesate (R. W. Taylor and J. Fee-
han).
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter samples, and in a rotting tree.
18.iii.1981 (W. L. Brown);
Pristomyrmex obesus Mann
Figures 233-236, 267, 278
Pristomyrmex obesus Mann, 1919; 339. Syntype
workers, queen and male, Solomon Is.: Ysabel, Fu-
lakora; Malaita, Auki; Three Sisters, Malapaina;
19.v.-24.xi.1916 (W. M. Mann) (MCZC, USNM,
AMNH) [examined].
Pristomyrmex pegasus Mann, 1919: 338. Holotype
worker, Solomon Is.: Santa Cruze: Graciosa Bay,
19.v.-24.xi1.1916 (W. M. Mann) (USNM) [exam-
ined]. Syn. n.
Pristomyrmex obesus subsp. melanoticus Mann, 1919:
340. Syntype workers, Solomon Is.: San Cristoval,
Pamua: Wai-ai; 19.v.—4.x1.1916 (W. M. Mann)
(MCZC, USNM) [examined]. Syn. n.
510
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 233-236. Pristomyrmex obesus Mann. 233: Worker head, full-face view; 234: Worker, lateral view; 235: Queen head,
full-face view; 236: Queen, lateral view.
Note: Pristomyrmex pegasus and P. obe-
sus were described as two new species in
Mann’s (1919) paper, on pages 338 and
339, respectively (i.e., P. pegasus appeared
before P. obesus; see also the previous ci-
tation). However, when proposing that
these two names are synonymic, I choose
P. obesus instead of P. pegasus as a valid
specific name for the following two rea-
sons: (1) Mann (1919) mentioned the pres-
ence of the elevated sides of the mesotho-
rax, and the absence of a median tooth on
the anterior clypeal margin are character-
istic of P. pegasus. However, “the elevated
sides of the mesothorax” are not shown in
the unique holotype of P. pegasus, and
“the absence of the median tooth on an-
terior border of clypeus” is actually an in-
dividual variation. (2) P. obesus possesses
about a dozen syntypes, including a female
and a male, but P. pegasus has only a single
type specimen (i.e., holotype). If the ho-
lotype of P. pegasus is lost or destroyed in
the future, it would be very inconvenient
to those people who want to see it. Thus,
P. obesus is selected as a valid name here.
Diagnosis (Worker). Masticatory margin
of mandible lacking a diastema and _ pos-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
sessing four teeth, of which the third
tooth, counting from the apex, smallest;
eyes containing = fou to seven ommatidia in
the longest row; dorsum of head, except
for the scrobes, usually with dense foveo-
late punctures; entire first gastral tergite
evenly covered with numerous erect or
suberect hairs.
Worker. TL 2.24-3.22, HL 0.63-0.86,
HW 0.61-0.86, CI 94-104, SL 0.52—0.75,
Sl 79-88, EL 0.09-0.14, PW 0.41—0.55,
AL 0.58—0.88, PPW 0.18—0.24, PPL 0.16—
020) PRL 105-125 Ga = 90).
Mandibles usually smooth and shining
but sometimes with a few superficial short
rugae or a few hair pits. A broad-based tri-
angular short tooth present about midway
on ahe basal margin of the mandible. Clyp-
eus flat, smooth, | aa the median carina of
frontal area usually extending to the clyp-
eus. Anterior clypeal margin ith a median
denticle and two lateral teeth but the me-
dian denticle often smaller than the oth-
ers; sometimes the median denticle lack-
ing; thus, only two teeth are present there.
Wentr al surface of clypeus lacking toothlike
prominences or rugae. Palp formal RSE
Frontal carinae extending to the level of
the posterior margins of eyes. Scrobal im-
pressions broad, Sellen: present lateral to
the frontal carinae. Frontal lobes slightly
expanded. Eyes moderately sized, usually
containing five to seven (rarely four) om-
matidia in the longest row. Profile of ali-
trunk and pedicel segments as in Figure
234. Pronotum unarmed. Propodeum with
a pair of triangular short spines. Meta-
pleur: al lobes rounded. Dorsum of alitrunk
in dorsal view usually with a longitudinal
impression or furrow at middle, bat some-
times this longitudinal impression indis-
tinct. Petiole mode in profile high with the
anterodorsal angle higher than nile poster-
odorsal, its Ankonor “peduncle nearly as
long as the node. Subpetiole with a narrow
semitranslucent lamella. Postpetiole in
profile rounded dorsally. In dorsal view,
dorsum of petiole node subrounded, about
as long as broad; postpetiole somewhat
transversally rectangular and _ slightly
511
broader than long. Dorsum of head, ex-
cept for the scrobes, usually with dense fo-
veolate punctures. Dorsum of alitrunk
with scattered foveolate punctures. Peti-
ole, postpetiole, and gaster smooth and
shining. Dorsal surfaces of head and ali-
trunk with numerous erect to suberect
long hairs. Two pairs of hairs usually pre-
sent on the dorsum of petiole node and
two to three pairs usually on the dorsum
of postpetiole. Entire first gastral tergite
covered with numerous, evenly distr cece
erect or suberect hairs. A few pairs of for-
ward-projecting hairs present near the an-
terior clypeal margin. Scapes and _ tibiae
with numerous erect to suberect short
hairs. Color reddish-brown.
Queen. TL 2.69-3.40, HL 0.69-0.82,
HW 0.67-0.85, CI 97-104, SL 0.54—0.68,
SIP 7S—Sose IE Onlds=0 199 RWaO54=0164
AL 0.76—0.94, PPW 0.20-0.26, PPL 0.18—
022) Peis 25 =s1'5)
Generally similar to worker, except for
caste differences. In addition, foveolate
punctures shallow on the mesonotum; pro-
podeal armaments slightly shorter than
those in the conspecific workers: dorsum
of alitrunk lacking a longitudinal furrow.
Male (Figs. 267, 278). A syntype male,
together with Alyontt a dozen workers and
a queen, collected in Fulakora, Solomen
Th. by W. M. Mann on 19.v.—24.xi.1916,
constitutes a series. A second male speci-
men, together with five workers, collected
in Guadalcanal, Solomen I., by P. Green-
slade, constitutes another series. Each of
the two males was originally mounted with
two or three workers, respectively, on the
same pin. TL 2.92; HL 0.52, 0.53; HW
03533) 0'54- CI 0250102 3S O22 0802235
38, 41: HWE 0.66, 0.68; EL 0.25, 0.26:
PW 0.58, 0.60; AL 0.90, 0.92; PPW 0.18;
PRE OS SPR LOOnGs—s2):
Head, including the eyes, broader than
long. Clypeus convex, without a median
longitudinal carina. Anterior clypeal mar-
gin transverse. Frontal carinae weak or in-
distinct. On the mesoscutum, notauli dis-
tinct, forming a Y shape; parapsidal fur-
rows absent. Scuto-scutellar sulcus rather
512
broad, with five narrow longitudinal ridg-
es. Propodeum weakly faberculate: lacking
teeth and spines. Metapleural lobes some-
what rounded. Petiole node in profile low,
rounded dorsally. Postpetiole in profile
low, rounded dorsally, in dorsal view sub-
quadrate. Dorsum of head unsculptured
and shining, except for a median longitu-
dinal carina present on the frontal area.
Dorsal surface of alitrunk smooth and
shining but with well-marked sutures. Pet-
iole, postpetiole, and gaster smooth and
shining. Dorsal surfaces of head, alitrunk
and gaster with abundant long hairs. Legs
ane numerous hairs. Body lacigehe
brown; wings infuscated.
Gonmmients: Pristomyrmex obesus occurs
on Solomon Is. It is similar in the workers
to P. simplex of New Guinea but can be
separated from the latter by possessing nu-
merous erect or suberect hairs evenly dis-
tributed on the entire first gaster tergite.
In the levigatus group, this character is
present only in two species, P. obesus and
P. longus; in the other species, the first gas-
tral tergite has a few or no hairs. In addi-
tion, foveolate punctures on the dorsal
head are denser in the workers of P. obesus
than in P. simplex.
Material Examined (ANIC, MCZC,
BMHH, USNM, BMNH#). Solomon Is.:
Ysabel, Fulakora (W. M. Mann); Guadal-
canal, Mt. Austen (P. Greenslade); Guad-
aleanal, Kukum (P. Greenslade); Guadal-
canal, Gold Ridge, 2,000 ft (P. Green-
slade); Guadaleanal, Mt. Jonapau, 2,600 to
3,500 ft (P. Greenslade); Guadalcanal, Na-
limbiu R. (P. Greenslade); Guadalcanal,
Balesuna R. (P. Greenslade); Guadalcanal,
Visale (P. Greenslade); Guadalcanal, Uma-
sani R., 1,000-ft ridge, leaf litter (P. N.
Lawtence); San Cristoval, Warahito R.,
275 ft (P. Greenslade); San Cristoval, for-
est, 250 ft (P. Greenslade); Malaita, Small
Malaita (P. Greenslade); Malaita, Dala (P.
Greenslade); New Georgia, Kolomban-
gara, Hunda (P. Greenslade): New Geor-
gia, Kolombangara, S. Kusi (P. Green-
slade); New Georgia, Kolombangara, N of
Kuzi, 500 ft, forest litter (P.N.L.); New
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Georgia, Vella Lavella, Bara koma (P|
Greenslade): New Georgia, Vangunu I. (P. |
Greenslade); Choiseul, Wagina I. (P
Greenslade); Russell Is., Luavic (P. Green-
slade); Nggela (P. Green lade). Isabel, Bu-
ala (P. Greenslade); Vanikord (P. Green- |
slade); Bougainville I., Panguna, 600 to.
800 m (J. i Gressitt). |
Ecological Information. This species has _
been eollemnadl | in forest litter. Mann (1919) |
found a colony composed of less than a
dozen workers, a dealated queen, and one |
male under a stone.
Pristomyrmex simplex sp. n.
Figures 237-240
Diagnosis (Worker). Masticatory margin |
of mandible lacking a diastema and pos-
sessing four ere of which the third
tooth, counting from the apex, smallest;
dorsal sinthioest of alitrunk and head bes
tween the frontal carinae with scattered
foveolate punctures; eyes usually contain-
ing five (rarely four) ommatidia in the lon-
gest row; a few hairs present on the first
gastral tergite.
Holotype Worker (ANIC). TL 2.48, HE
0.66, HW 0.69, CI 105, SL 0.52, SI 7&
EL 0.12, PW 0.46, AL 0.58. Paratypes, six
workers and three queens (MCZC, ANIC,
BMNH).
Worker. TL 2.42-2.64, HL 0.65—0.683
HW 0.66-—0.70, CI 100—105, SL 0.52—0.54,
SI 74-79, EL 0.09-0.12, PW 0.44—0.46;
AL 0.58—0.66, PPW 0.18—0.20, PPL 0.14—
0.16, PPI 119-129 (n = 6).
Mandibles usually smooth and shining
but sometimes with a few small shallow
hair pits and a few superficial short rugae.
A broad-based triangular short tooth pre-
sent about midway on the basal margin of
the mandible. Clypeus flat, emoothe and
shining, usually unsculptured but some-
times fie median carina of the frontal area
extending to the clypeus. Anterior clypeal
margin wath a median denticle and two lat-
eral teeth; the median denticle is often
smaller than the others; sometimes the
median denticle is absent so that the only
two teeth are present there. Ventral sur-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
513
Figures 237-240. Pristomyrmex simplex sp. n. 237: Worker head, full-face view; 238: Worker, lateral view; 239: Queen head,
full-face view; 240: Queen, lateral view.
face of clypeus lacking rugae or toothlike
prominences. Palp formula 1,3. Frontal ca-
rinae distinct, extending to the level of the
posterior margins of eyes. Scrobal areas
slightly concave, present lateral to the
frontal carinae. Frontal lobes weakly ex-
panded. Eyes moderately sized, about 0.14
to 0.17 X HW, usually containing five
(sometimes four) ommatidia in the longest
row. Occipital margin feebly concave in
full-face view. Profile of alitrunk and ped-
icel segments as in Figure 238. Pronotum
unarmed. Propodeum with a pair of tri-
angular teeth. Metapleural lobes rounded.
Petiole node in profile high with the an-
terodorsal angle higher than the postero-
dorsal, its anterior peduncle about as long
as the node. Subpetiole with a narrow rim.
Postpetiole in profile rounded dorsally. In
dorsal view, dorsum of petiole node tran-
soval and dorsum of postpetiole somewhat
transversely rectangular. Dorsum of head
between the frontal carinae with scattered
foveolate punctures, varying from a few
feeble punctures to numerous distinct
ones; spaces between foveolae often
514
smooth. Dorsum of alitrunk with a few to
some scattered foveolate punctures. Peti-
ole, postpetiole, and gaster smooth and
shining. Dorsal surfaces of head and ali-
trunk with numerous erect to suberect
hairs. Two pairs of hairs usually present on
the dorsal surfaces of petiole node and
postpetiole, respectively, and a few near
the base of the first gastral tergite. A few
pairs of forward-projecting hairs present
near the anterior clypeal margin. Scapes
and tibiae with some erect to suberect
short hairs. Color reddish-brown.
Queen. TL 2.51—2.84, HL 0.66—0.70,
HW 0.66—0.71, CI 1OO—101, SL 0.51—0.58,
SI 76-82, EL 0.14—0.16, PW 0.50-0.55,
AlevO166-0:S81, BRW.0N9=0015 PRM ON5=
ONS) PPI 114=1335@7. = 3):
Generally similar to worker, except for
caste differences. In addition, mesoscutum
rather smooth and shining, but mesoscu-
tellum with a few foveolate punctures;
propodeal armaments slightly shorter than
those in the conspecific worker.
Male. Unknown.
Comments. This species occurs in New
Guinea. It is closely related to P. levigatus
and P. obesus. It is separable from P. levi-
gatus by possessing some foveolate punc-
tures on the dorsum of the alitrunk in the
workers and on the dorsal head between
the frontal carinae in the workers and
queens. It differs from P. obesus because
the workers of P. simplex have only a few
hairs on the first gastral tergite. Pristomyr-
mex sinylex differs from P. coggii by the
following characters: The eyes usually con-
tain five (sometimes four) ommatidia in
the longest row in the workers of P. sim-
plex but two to three ommatidia in P. cog-
gii; the dorsum of the head has foveolate-
reticulate sculpture in the workers and
queens of P. coggii but scattered foveolate
punctures in P. simplex.
Holotype. Papua: 8 km S of Kokoda, S00
m, rainforest, ANIC Berleasate, No. 382,
L.vi.1972 (R. W. Taylor).
Paratypes. One worker with same data
as holotype; two workers, Papua: Karema,
Brown R., lowl. rainfor., 8—11.iii.1955 (E.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
O. Wilson); three workers and one queen,
Papua New Guinea: Port Moresby, Brown |
River, 2.x.1969 (J. Balogh); one queen;
Papua: N. Dist., 27.xii.1971 (P. M. Room);
one queen, Papua: N. Dist., Debelou,
23.vi.1973 (P. M. Room).
Non-Type Material Examined. A worker
(NHMV), collected in the Philippines (Lu-_
zon, Lagunas, Mt. Banahaw above Kina-
buhayan, 600-700 m) by J. Kodada and B.
Rigova, has the following measurements:
TL 2.98, HL 0.84, HW 0.84, CI 100, SE
0.70, SI 83, EL 0.10, PW 0.54, AL 0.748
A second worker (LAMN), collected in
New Guinea (Gulf Prov., Ivimka Camp,
Lakekamu Basin, 7.73°S, 146.76°E, 120 m,
#96-291, lowland wet forest, ex sifted leaf
litter) by R. R. Snelling, bears a few small,
feeble, shallow punctures on the dorsal al-
itrunk; this specimen possesses TL 2.48,
HL 0.68, HW 0.69, CI 101, SL 0.58, Sil
84, EL 0.09, PW 0.45, AL 0.62.
Ecological Information. This species oc-
curs in rainforest and has been collected
in litter samples.
THE PROFUNDUS GROUP
Worker. Small sized, with the following
combination of characters.
(1) Masticatory margin of mandible
lacking a diastema and possessing four
teeth (i.e., the strongest apical + the sec-
ond strongest preapical + the smallest
third + an acute basal tooth); basal margin
of mandible with a strongly prominent
tooth that is adjacent to the basal tooth of
the masticatory margin; as a result, five
teeth are set close together.
(2) Lateral portions of clypeus in front
of antennal fossae reduced to margins.
(3) Antennal scrobes broad, deep, and
extending close to the occipital corners.
(4) Base of antennal scape lacking a cir-
cling lamella.
(5) Mesonotum much higher than pro-
podeal dorsum so that the dorsum of ali-
trunk in profile is not continuously arched.
(6) Metapleural lobes vestigial and in-
distinct.
The profundus group is closely related
to the levigatus group. The form of den-
tition of the masticatory margin of the
mandible is a critical character shared by
the workers and queens of the two groups.
Possessing so many autapomorphie char-
acters, including that a tooth on the basal
margin of the mandible is adjacent to the
basal tooth of the masticatory margin, as
well as the previously mentioned charac-
ters 3 to 6, the profundus group is easily
separable from the levigatus group and all
other Pristomyrmex species groups.
This group contains only a single spe-
cies, P. profundus of Malaysia.
Pristomyrmex profundus sp. n.
Figures 241-244
Diagnosis (Worker). See characters 1
and 5 melee the profundus group.
Holotype Worker (BMNH). TL 2.86,
HL 0.64, HW 0.74, CI 119, SL 0.44, SI
59, EL 0.13, PW 0.52, AL 0.68, PPW 0.24,
PPL 0.16, PPI 150. Paratypes, 19 workers
and three queens (BMNH, MCZC,
MHNG, ANIC, LAMN).
Worker. TL 1.74—2.86, HL 0.46-0.66,
HW 0.46-0.74, CI 96-119, SL 0.36-0.44,
Slo 97, 9 Ee OL09=0545 RW. 0!34=0:522
mls O.44=0.68: PEW O115-0.24 PPL 0.11—
lo. PRL 123-645 m= 20):
Mandibles usually smooth and shining
but sometimes with a few weak, short, bas-
al rugae. Clypeus not depressed, usually
with a median longitudinal carina and two
transverse carinae. Anterior clypeal margin
lacking any distinct denticles. Ventral sur-
face of clypeus lacking any toothlike prom-
inences but usually with a weak transverse
ruga. Palp formula undissected. Frontal
carinae divergent, extending beyond the
level of the posterior margins of eyes and
close to the occipital margin. Frontal lobes
completely absent. Antennal scrobes deep,
wide, present between the frontal carinae
and the eyes for the reception of the
scapes and funiculi of antennae. Antennal
scapes, when lying in the scrobes, close to
the occipital margin of head. Eyes contain-
ing five to eight ommatidia in the longest
row. In full-face view, head widest near the
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
515
occipital corners; occipital margin rather
straight. Dorsum of alitrunk in profile not
continuously arched, with the mesonotum
much higher than the propodeal dorsum
(i.e., a vertical cliff present between the
mesonotum and the propodeal dorsum).
Pro-mesonotum forming a single convex.
Sometimes the mesonotum higher than
the pronotum; thus, in dorsal view, a pro-
mesonotal impression is seen. Pronotum
unarmed. Propodeum with a pair of mod-
erately long spines. Metapleural lobes ves-
tigial and indistinct. In profile view, petiole
node high, ca. 0.21 to 0.29, much higher
than long and also higher than postpetiole,
with a single evenly blunt-rounded apex
and a long anterior peduncle. Subpetiole
with a narrow lamella. Postpetiole in pro-
file high, ca. 0.15 to 0.23, distinctly higher
than long, with a rounded dorsum. In dor-
sal view, postpetiole transoval and much
broader than long. Dorsum of head, ex-
Cope tOmathe atennl scrobes, fully cov-
ered with rugoreticulum. Galen sculpture
present on tie pro-mesonotum. Propodeal
dorsum with a few longitudinal rugae. Pet-
iole smooth and shining, usually with a
longitudinal carina on Sch side. Postpe-
Bole and gaster smooth and shining. Dor-
sal sirtaces of head, alitrunk, aide gaster
with numerous erect, thick, long eae
sometimes some hairs somewhat clavate A
pair of hairs present on the dorsum of pet-
iole node and usually two pairs on the dor-
sum of postpetiole. A few pairs of forward-
projecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect or suberect moderately long
hairs. Color light yellow to yellow brown.
Queen. PIL, 3 46—5592 ee Euley OL62=0:635
HW 0.76—-0.77, CI 123-124, SL 0.44—-0.46,
Si o7— Ole EL Onl5=0s638 RW 710'66=0570)
AL 0.92-0.98, PPW 0.28—0.29; PPL 0.18—
(0) A0), IAP TAO (= S&S).
General shape as in Figures 243-244,
with normal caste differences from the
conspecific worker; mesonotum unsculp-
tured, smooth, and shining; anterior end
of mesoscutum medially slightly concave.
Other characters similar to worker.
516
P FREES =~
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 241-244. Pristomyrmex profundus sp. n. 241: Worker head, full-face view; 242: Worker, lateral view; 243: Queen head, |
full-face view; 244: Queen, lateral view.
Male. Unknown.
Comments. This is a distinct species be-
cause many characters possessed by its
workers and queens are unique in the ge-
nus. In the entire Pristomyrmex fauna,
three species (P. profundus, P. divisus, and
P. pulcher) do not possess any denticles on
the anterior clypeal margin, but the latter
two species belong to the punctatus group.
Holotype Worker. Malaysia: Sabah, Por-
ing Hot Springs, 500 m, 7.v.1987 (Burck-
hardt and Lobl).
Paratypes. Seventeen workers and three
queens with same data as holotype; two
workers with same data as holotype but
date 6.v.1987.
Ecological Information. Unknown.
THE UMBRIPENNIS GROUP
Worker. Medium to large sized (HL |
1.04-1.68, HW 1.02-1.74, TL 4.10-7.06)
with the following combination of charac-
ters.
(1) Masticatory margin of mandibles _
with four teeth arranged as the strongest.
apical + the second strongest preapical +
two broad-based short teeth of similar size;
diastema lacking or indistinct.
(2) Basal margin of mandible with a)
central, broad-based, prominent lobe.
(3) Lateral portions of clypeus in front |
of antennal fossae reduced to a margin.
(4) Anterior margin of the median por-
tion of clypeus usually with five to seven
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
517
246
Figures 245-246. Pristomyrmex fuscipennis (F. Smith). 245A: Queen head, full-face view; 245B: Showing a transverse carina
on the ventral clypeus; 246: Queen, lateral view.
denticles; lateral portions of anterior clyp-
eal margin in front of antennal fossae with
a few weak blunt denticles.
(5) Ventral surface of clypeus with a
transverse ridge.
(6) Frontal lobes present, partly cover-
ing the condylar bulbs of holding antennal
scapes.
(7) Frontal carinae extending to the lev-
el of the posterior margins of eyes.
(S) Lamella that encircles the base of
antennal scape usually with a broad and
deep notch on the center of the dorsal sur-
face.
(9) Palp formula 1,3.
(10) Eyes small; EL is about 0.040 to
0.064 X HW in P. picteti and P. pollux and
0.069 to 0.108 k HW in P. umbripennis.
(11) Alitrunk in profile, not including
propodeal spines, with a regularly arched
dorsum, in dorsal view without any su-
tures.
(12) Pronotum unarmed.
(13) Metapleural lobes bluntly round-
ed.
(14) Petiole node in profile longer than
high, with a long anterior peduncle; its an-
cenadlonsall angle is on approximately the
same level as the posterodorsal.
(15) Foveolate punctures or foveolate-
reticulate sculpture present on the dorsal
surfaces of the head and the alitrunk.
This is a monophyletic group, contain-
ing five valid species. It is easily recogniz-
able by possessing characters 1, 2,5, 8, and
14. This group is endemic in the Oriental
region and restricted to the Philippines,
Malaya, Singapore, Brunei, Sabah, Bor-
neo, Indonesia, and Papua New Guinea,
The males of the wumbripennis group are
easily distinguished from the oer known
Pristomyrmex males by possessing the fol-
lowing characters: (1) medium to large
size; (2) palp formula 1,3; (3) propodeum
with a pair of broad-based, robust spines
(Figs. 256, 279, 280); and (4) the sides of
petiole with some longitudinal or reticu-
late rugae (Figs. 256, 279, 280).
Pristomyrmex fuscipennis (F. Smith)
Figures 245-246
Myrmica fuscipennis F. Smith, 1861: 46. Holotype
a Indonesia: Celebes, Tondano (A. R. Wal-
lace) (OXUM) [examined].
? Pristomyrmex fuscipennis (F. Smith) Emery, 1901:
567.
Pristomyrmex fuscipennis (F. Smith)
1932: 468.
Donisthorpe,
Queen. TL 6.92, HL 1.62, HW 1.64, Cl
OM SE MEAG Sl -SOn Hl 032 ee ENVIR S 2D:
AL 1.98 (n = 1).
Comments and Discussion. This species,
described from a single queen, obviously
belongs to the wmbripennis group by pos-
sessing the following characters: (1) mas-
ticatory margin of Sanandible with four
518
teeth (an apical + a preapical + two
broad-based short teeth of similar size),
lacking a distinct diastema; (2) basal mar-
gin of mandible with a central, broad-
based, prominent lobe; (3) frontal lobes
partially covering the condylar bulbs of
holding antennal scapes; (4) lamella that
encircles the base of antennal scape with
a broad and deep notch on the center of
the dorsal surface; (5) a coarse transverse
carina present on the ventral surface of
clypeus; and (6) anterodorsal angle of pet-
iole node in profile on approximately the
same level as the posterodorsal.
This queen differs from the queens of
P. picteti, P. pollux, and P. umbripennis as
follows: In P. fuscipennis, the dorsum of
the head possesses foveolate-reticulate
sculpture; many foveolate punctures be-
tween the frontal carinae are almost
aligned so that it seems that several lon-
gitudinal rugae are present there. These
longitudinal rugae are indistinct or absent
in P picteti, P. pollux, and P. umbripennis.
In P. picteti and P. umbripennis, the dor-
sum of the head possesses scattered fove-
olate punctures. In addition, in P. fusci-
pennis, the antennal scapes lack longitu-
dinal carinae along their dorsal margins, a
median longitudinal carina runs through
the entire clypeus and frontal area, dad
only five teeth are present on the anterior
margin of the median portion of the clyp-
eus, Sich are different from those in P.
pollux.
Pristomyrmex fuscipennis may be a sib-
ling species of P. picteti because the queen
of P. fuscipennis is very similar to that of
P. picteti, except for possessing foveolate-
reticulate sculpture and longitudinal rugae
on the dorsal head. In Battin: I have ex-
amined several workers that may belong to
P. fuscipennis. The cephalic sculpture of
these workers is similar to that of the
queen of P. fuscipennis, but the other
characters of the workers are identical
with those of the workers of P. picteti.
Finally, I feel that P. fuscipennis may
represent an incipient species. Further
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
collecting and ecological investigations are
needed.
Ecological Information. Unknown.
Pristomyrmex picteti Emery
Figures 247-250, 268, 279
Pristomyrmex picteti Emery, 1893: 190. Lectotype
worker, Sumatra: Deli (Bedot) (MCSN), here des-
ignated, [examined].
Pristomyrmex picteti var. tingiana Stitz, 1925: 120.
Syntype workers, Philippines: N. Palawan, Binal-
uan, xi-xii 1913 (G. Boettcher) (MNHU, MCZC)
[examined]. Syn. n.
Diagnosis (Worker). Masticatory margin
of mandible with four teeth (an apical + a
preapical + two broad-based short teeth
of similar size), lacking a distinct diastema;
eyes small, with three to four ommatidia
in the longest row; propodeal spines fairly
long, about 1.3 to 1.5 x the distance be-
tween their bases, not strongly upcurved
at their apices.
Worker ls 4110=5:84-) lge L04= 1836"
HW 1.02—1.40, CI 96—108, SL 0.92-1.26;
SI 85-97, EL 0.05—0.09, PW 0.68—0.88,
AL 1.12—1.58, PPW 0.30-0.41, PPL 0.274
0:36, PRE 100=120°@-=40))
Mandibles rather smooth or sometimes
with a few longitudinal rugae. Basal mar-
gin of Tanchible with a br eagle based, sub-
triangular, short prominence or a some-
eohate curved lobe. Clypeus smooth and
shining, except for a median longitudinal
carina hat usually crosses the entire clyp-
eus but sometimes does not reach the pos-
terior clypeal margin. Anterior margin of
the median portion of clypeus (not ind
ing the margins in front of the antennal
fossae) with a median denticle and two
(sometimes three) lateral denticles on each
side. Ventral surface of clypeus with a
coarse, transverse, long carina. Palp for-
mula 1,3. Frontal carinae usually beyond
the eyes. Antennal scrobes indistinct.
Frontal lobes present, partially covering
the condylar bulbs of holding antennal
scapes. Antennal scapes, when lying on the
dorsal head, just reaching or slightly be-
yond the occipital margin. Lamella, encir
cling the base of antennal scape, usually
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
519
Figures 247-250. Pristomyrmex picteti Emery. 247A: Worker head, full-face view; 247B: Showing a transverse carina on the
ventral clypeus; 248: Worker, lateral view; 249A: Queen head, full-face view; 249B: Showing a long transverse carina on the
ventral clypeus; 250: Queen, lateral view.
with a broad and deep notch on the center
of the dorsal surface, but in several spec-
imens (from Palawan I., Philippines), this
lamella is entire, without a notch. Eyes
very small, usually with three, at most four,
ommatidia in the longest row. Occipital
margin straight or very feebly concave in
full-face view. Alitrunk in profile with a
convex dorsum. Pronotum unarmed. Pro-
podeum with a pair of fairly long spines,
ca. 1.3 to 1.5 X the distance between their
bases; sometimes the spines are weakly
upcurved at their apices. Metapleural
lobes rounded or somewhat truncated.
Petiole node in profile slightly longer than
high, with a fairly long anterior peduncle;
its anterodorsal angle is usually on approx-
imately the same level as the posterodor-
sal, but sometimes the former is slightly
higher than the latter. Subpetiole with a
narrow, semitranslucent lamella. Postpeti-
ole in profile higher than long, rounded
dorsally, in dorsal view broadening from
front to back. Dorsal and ventral surfaces
of head, dorsum of alitrunk, as well as the
sides of pronotum with numerous foveo-
late punctures; space between foveolae of-
ten smooth. Antennal scapes smooth, or
with one to two longitudinal rugae along
their margins. Petiole smooth and shining,
except for a longitudinal ruga on each side.
Postpetiole and gaster smooth and shining.
520
Dorsal surfaces of head and alitrunk with
numerous erect or suberect short hairs.
Two or more pairs of hairs present on the
dorsal surfaces of petiole node and post-
petiole, respectively. A few of forward-pro-
jecting long hairs present near the anterior
clypeal margin. Scapes and tibiae with nu-
merous erect or suberect short hairs. First
gastral tergite lacking erect or suberect
Raine Color reddish- Broun
Queen. TL 6220=72025) Ei e264"
HW 1.42=1.69) CI 103-1138, SL 1.21—1.56,
SI 85-92, EL 0.29-0.34, PW 1.14-1.42,
Alene (4=2 1A PRWa041= 0525 PRI 036=
0.42, PPI 110-125 (n = 13).
General shape as in Figures 249-250,
with normal caste differences from the
conspecific worker; eyes much larger, usu-
ally containing more than 17 onuneuchan in
the longest row; other characters similar to
worker.
Male (Figs. 268, 279) la AG=> 2 Se Ee
0.68—0.78, HW 0.69—0.80, CI 95-108, SL
0.30—0.42, SI 40-56, HWE 0.85—0.94, EL
0.35-0.39, PW 0.92-1.14; AL 1.44—-1.78,
PPW 0.30-0.38, PPL 0.28—0.34, PPI 103—
114 (n = 5).
Head in full-face view, including the
eyes, broader than long. Clypeus convex,
lacking a median longitudinal carina. Palp
formula 1,3. ronal carinae distinct, ex-
tending to the level of the posterior mar-
gins of antennal insertions. Maximum di-
ameter of the median ocellus 0.10 to 0.12.
Scapes longer than the other antennal seg-
ments, except for the apical ones. On the
mesoscutum, notauli rather wide, forming
a V shape, separated into several cells by
narrow transverse ridges; parapsidal fur-
rows absent. Scuto-scutellar sulcus wide,
usually with five to six narrow longitudinal
ridges. Propodeum armed with a pair of
robust, triangular short spines. Metapleur-
al lobes subtriangular. Middle and hind
tibiae without any spurs. Petiole node in
profile low, slightly longer than high, with
a fairly long anterior peduncle. Postpetiole
in profile rounded dorsally, in dorsal view
somewhat transversely rectangular and
slightly broader than long. Dorsum of
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
head with some small punctures, varying
from scattered and shallow to dense and
somewhat coarsely incised. Pronotum with —
dense foveolate punctures. Mesonotum |
with punctures, varying from a few scat-
tered to numerous and from feeble, small
to rather large. Propodeum rugulose. Sides
of petiole with a few longitudinal rugae as
well as some foveolate punctures between
them. Sides of postpetiole with some shal-
low foveolate punctures. Gaster unsculp-
tured. All dorsal surfaces with numerous
blackish-brown long hairs; sometimes hairs _
somewhat stiff. Color blackish-brown;
wings dusky.
Note: The previously described male is
assigned to the species P. picteti for the
following three reasons: (1) It belongs to
the umbripennis group because it is very
similar to the males of P. pollux, P. reti-
culatus, and P. umbripennis but rather dif-
ferent from the other known males of Pris-
tomyrmex in the structure and shape of
propodeal spines, petiole, notauli, and scu-
to-scutellar sulcus and in body size, sculp-
ture, and hairs. (2) It differs from the
males of the other species of the wmbri-
pennis group as follows: In the male of P.
picteti, CI is 95-108, EL is 0.35—0.39, the
propodeal spines are shorter, the postpe-
tiole in dorsal view is slightly broader than
long (PPI 103-114), and the notauli form
a V shape; but in P. pollux, CI is 80, EL
is 0.44, the propodeal spines are longer,
the postpetiole is distinctly longer than
broad (PPI 80-90), and the notauli form a
Y shape. The male of P. picteti (HW 0.69—
0.80, HL 0.68—0.78, and EL 0.35-—0.39) is
smaller than that of P. umbripennis (HW
0.98, HL 0.94, and EL 0.47). A distinct
rugoreticulum is seen on the dorsal head
and also on the mesonotum in P. reticu-
latus but not so in the male of P. picteti.
(3) Two male specimens were collected in
Dumaguete, Philippines, by J. W. Chap-
man on May 30, 1950, and each of them
was originally mounted together with one
worker or one queen of P. picteti, respec-
tively, on the same pin.
Comments. Pristomyrmex picteti occurs
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
in Papua New Guinea, Borneo, Sabah,
Brunei, Malaya, Singapore, and the Phil-
ippines. It is closely related to the other
several species of the wmbripennis group.
A discussion regarding the relationship be-
tween P. picteti and P. fuscipennis is pro-
vided under P. fuscipennis. Characters
separating P. picteti from P. umbripennis
are listed under P. umbripennis. The work-
ers of P. picteti differ from those of P. pol-
lux as follows:
P. picteti
Propodeal spines relatively short, not
strongly upcurved at their apices
Smaller species with HL 1.04—1.36, HW
1O2= "40" andi Sie 0:92=1-26
Dorsum of head with scattered foveo-
late punctures; space between foveo-
lae often smooth
Anterior margin of the median portion
of clypeus usually with fewer than
seven denticles
P. pollux
Propodeal spines relatively long, strong-
ly upcurved at their apices
Larger species with HL 1.42-1.54, HW
Zea sey emo! Si, 40S ia
Dorsum of head with foveolate-reticu-
late sculpture
Anterior margin of the median portion
of clypeus usually with seven denti-
cles
Material Examined (ANIC, BMHH,
BMNH, LAMN, MCZC, NHMV). Papua
New Guinea: Gulf Prov., Ivimka Camp,
Lakekamu Basin, 7.7°S, 146.8°E, 120 m el-
-evation, lowland wet forest, #96-395 (nest
in wet rotten log), #96-404 (in rotted log
debris) (R. R. Snelling); NETH. Genjam,
40 km W of Hollandia, 100 to 200 m (T.
C. Maa); 24 km N Madang, 80 m, 5°01'S,
145°46'E, rainforest, ex rotten log (P. S.
Ward); Yawasora near Wewak, <50 m,
rainforest, ex rotting log (R. W. Taylor);
along Kokoda Rd., rainforest, 400 to 1,000
ft (B. B. Lowery); Lae, Didiman CK., low-
land rainforest (E. O. Wilson); Lae, Busu
R., lowland rainforest, in rotten wood on
ground (B. B. Lowery); Lae, Markham R.
521
Bridge, lowland rainforest, in log (B. B.
Lowery); Lae 300 ft, above Bumbu Crk.,
edge of rainforest, wet gully, in rotten
wood (B. B. Lowery); Tage L. Kutubu
2700 tte forest maxreiny ext leavesn (pl:
Barrett); Papua, Brown Riv., lowland rain-
forest, ex rotten wood fragment (R. W.
Taylor). Indonesia: Irian Jaya, PT. Free-
port Concession, Siewa camp., 03.04°S,
136.38°E, 200 ft, #98-86, lowland second-
ary rainforest, under loose bark of dead log
(R. R. Snelling); Borneo, West Kaliman-
tan, Gunung Palung Nat. Pk. Cabang Panti
Ress Sta 00 sto 400 mmnee el S4SaellOa ke
primary forest (Datling, Rosichon, Sutris-
no): Borneo, Sandakan (Baker). N. Bor-
neo: Tawau, Quoin Hill, Forest Camp 1, 3
to 5 km WSW of Cocoa Res. Sta. (Y. Hir-
ashima). Brunei: Belait District, Manilas,
Ulu Belait (D. E. Hardy). Malaysia: Ma-
laya, Gombak (B. Bolton); Pahang, Kuala
Tahan 200 m (J. L. Gressitt); Sg. Patani (G.
H. Lowe). Singapore (Baker). Philippines:
Mt. Montalban, Rizal Wa-wa Dam, 150 to
200 m (H. M. Torrevillas); Mt. Apo 5 to
6,000 m (C. F. Clagg); N. Palawan, Bin-
aluan (G. Boettcher); Palawan, 14 km S
Puerto Princesa (9.44°N, 118.44°E), rain-
forest 0 to 250 m (B. B. Lowery); Palawan,
Iwahig Penal Col., ca. Puerto Pricesa
(9.44°N, 118.44°E), coffee plantation, 60
m, edge of rainforest, foraging on ground
(B. B. Lowery); Luzon, Los Banos, Mt.
Makiling 600 m (14.1°N, 121.11°E), rain-
forest, in very wet log (B. B. Lowery); Los
Banos (F. X. Williams); Mt. Makiling (Bak-
er; A. T. Cencho); Laguna, Mt. Makiling,
150 to 500 m (H. Zettel); Luzon I., Bauqui
(R.C.Mcg.); Luzon, Mt. Banahao (?); Du-
maguete, 4,500 ft (J. W. Chapman); Horns
of Negros, 3,600 ft (J. W. Chapman);
Camp (J. W. Chapman).
Ecological Information. This species oc-
curs in rainforest, nesting in rotten logs. It
has been collected on the ground, on rot-
ten wood, and under loose bark of a log.
Pristomyrmex pollux Donisthorpe
Figures 251-254, 269, 280, 282
Pristomyrmex pollux Donisthorpe, 1944: 83. Holo-
type male, Malaysia: Penang, 6.xi.1913 (G. E. Bry-
ant) (BMNH) [examined].
Figures 251-254. Pristomyrmex pollux Donisthorpe. 251A: Worker head, full-face view; 251B: Showing a transverse carina on
the ventral clypeus; 252: Worker, lateral view; 253A: Queen head, full-face view; 253B: Showing a transverse ruga on the ventral
clypeus; 254: Queen, lateral view.
Note: This species was described only
from a single male, which created the dif-
ficulty of associating female castes with the
male. I place the following workers under
P. pollux for three reasons: (1) In MCZC,
two males of P. pollux appear to belong to
the same series as the following examined
28 workers bearing labels with the same
locality and collector and with the absence
of collecting date. (2) This male (i.e., the
holotype of P. pollux) belongs to the um-
bripennis group because it is very similar
to the males of P. picteti, P. reticulatus, and
P. umbripennis but rather different from
the other known males of Pristomyrmex in
the structure and shape of propodeal
spines, petiole, notauli, and scuto-scutellar
sulcus and in body size, sculpture, and
hairs. (3) After all available males of Pris-
tomyrmex are examined, it seems that, in
Pristomyrmex, the propodeal armaments
of the male are usually shorter than those
of the conspecific worker. The male of P.
pollux has a pair of well-developed pro-
podeal spines (which are actually the
strongest and longest among all known
Pristomyrmex males). The following work-
ers also possess a pair of long propodeal
spines, which matches with those in the
male of P. pollux closely.
Diagnosis (Worker). Masticatory margin
of mandible with four teeth (an apical + a
preapical + two broad-based short teeth
of similar size), lacking a distinct diastema;
eyes small, with three to four ommatidia
in the longest row; propodeal spines long,
strongly upcurved at their apices.
Worker. TL 6.26—-6.80, HL 1.42—1.54,
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
HW 1.42-1.58, CI 95-104, SL 1.40-1.52,
SI 96-101, EL 0.06—0.08, PW 0.90—0.96,
AL 1.58—1.90, PPW 0.39-0.43, PPL 0.39—
0:43, PPI 100105 (7 = 28).
Mandibles with a few longitudinal ru-
gae. Basal margin of ToEtRCHS with a
roel short, soe wuhiat truncated, promi-
nent lobe. Clypeus with a median longi-
tudinal carina that does not reach the pos-
terior clypeal margin. Frontal area un-
sculptured. Anterior margin of the median
portion of clypeus with a median denticle
and three (sometimes two) others on each
side. Ventral surface of clypeus with a
coarse transverse carina. Palp formula 1,3.
Frontal carinae strong, extending to the
level of the posterior margins of eyes.
Scrobal areas smooth, present lateral to
the frontal carinae. Frontal lobes present,
partially covering the condylar bulbs of
holding Antennal scapes. Antennal scapes,
when lying on the dorsal head, just beyond
the occipital margin. Lamella, encircling
the base of antennal scape, with a broad
and deep notch on the center of the dorsal
surface. Eyes small, usually with three, at
most four, ommatidia in the longest row.
Occipital margin straight or elke: concave
in full-face view. Bronouiin unarmed. Pro-
podeum with a pair of long spines that are
strongly upcurved at their apices (i.e.,
hook Eee) and laterally compressed. Meta-
pleural lobes prominent and somewhat
rounded. Petiole node in profile distinctly
longer than high, with a long anterior pe-
Aamele: its Al terosloreal angle is on ap-
proximately the same level as the poster-
odorsal. In dorsal view, petiole node longer
than broad. Subpetiole with a narrow,
semitranslucent lamella. Postpetiole in
pr ofile higher than long, rounded dorsally,
in dorsal view broadening from front to
back. Dorsum of head, except for scrobal
areas and frontal area, with foveolate-re-
ticulate sculpture. Antennal scapes with
longitudinal rugae along their dorsal mar-
gins. Dorsum ae alntiannk as well as the
sides of pronotum with numerous foveo-
late punctures that are often close to each
other. Sides of the rest of alitrunk with
523
some scattered punctures. Petiole smooth
and shining, but with a longitudinal ruga
on each side. Postpetiole and gaster
smooth and shining. Dorsal surfaces of
head, alitrunk, petiole, and postpetiole
with numerous erect or suberect hairs. A
row of forward-projecting long hairs pre-
sent near the anterior clypeal margin.
Scapes and tibiae with some erect or sub-
erect short hairs. First gastral tergite lack-
ing erect or suberect hairs. Color yellow-
brown to reddish-brown.
Queen. A single queen (BMNH) was ex-
amined. It was collected in Penang, Ma-
laysia, by G. E. Bryant in October 1913
(ie., the collecting locality and the collec-
tor name for this specimen are the same
as the holotype male of P. pollux): TL ca.
(oO Ely 164 AW arco s Close Siw a6:
SI 92, EL 0.29, PW 1.42, AL 2.14.
This queen possesses the following
characters: (1) dentition of the masticatory
margin and basal margin of mandible as in
the previously Hcntoncd worker; (2) den-
tition of the anterior clypeal margin as in
the previously mentioned worker; (3 ) clyp-
eus with a median carina that does not
reach the posterior clypeal margin; (4)
ventral surface of clypeus with a transverse
ruga; (5) frontal lobes and frontal carinae
as in the previously mentioned worker; (6)
lamella encircling the base of antennal
scape as in the previously mentioned
worker; (7) antennal scapes with longitu-
dinal rugae along their dorsal margins; (8)
propodeum with a pair of robust, rather
long spines that are longer than the dis-
tance between their bases. (9) metapleural
cee as in the previously mentioned work-
; (10) petiole and postpetiole as in the
Saree mentioned worker; and (11)
dorsum of head, except for the scrobal ar-
eas, with foveolate-reticulate sculpture. In
other words, except for normal caste dif-
ferences and the propodeal spines that are
neither laterally compressed nor upcurved
at their apices, other characters are gen-
erally similar to those in the previously
mentioned worker.
Male (Figs. 269, 280, 282), Th 5.60=
524
5.92, HL 0.85-0.88, HW 0.68—-0.70, CI
80-80, SL 0.38-0.41, SI 54-60, HWE
0.86—-0.88, EL 0.44—0.44, PW 1.26—1.36,
AL 2.00—2.20, PPW 0.35—0.38, PPL 0.42—
0.44, PPI 80-90 (@ = 3).
Head in profile high and thick, in full-
face view, excluding eyes, much longer
than broad, and including the eyes about
as long as_ broad. Clypeus convex and
ar chee in middle without a median longi-
tudinal carina. Palp formula 1,3; maxillary
palp long. Maximum diameter of the me-
dian ocellus 0.12. Scapes longer than the
other antennal segments, except for the
apical ones. On the mesonotum, notauli
rather wide and deep, showing a Y shape,
separated into small cells by narrow trans-
verse ridges; parapsidal furrows absent.
Scuto- soutellar sulcus wide, separated into
small cells by seven to eight narrow ridges.
Propodeum ‘with a pair of robust, broad-
based, rather long spines. Metapleural
lobes prominent and somewhat rounded.
Petiole node in profile distinctly longer
than high with a long anterior peduncle.
Postpetiole in profile “slightly longer than
high with a convex dorsum, in don sal view
rectangular and distinctly longer than
broad. ‘Subpostpetiole with a blunt, tooth-
like prominence. Dorsum of head cath nu-
merous small foveolate punctures that
sometimes are coarse and dense. Prono-
tum with small, coarse, and dense foveo-
late punctures. Mesoscutum with some
scattered, small, shallow foveolate punc-
tures. Mesoscutellum with dense, coarse
foveolate punctures. Propodeum with
some irregular coarse rugae. Middle and
hind tibiae without any spurs. Each side of
petiole with a coarse longitudinal ruga, a
few irregular short rugae, and some foc
olate punctures. Side ae postpetiole with
some small, weak foveolate punctures. All
dorsal surfaces with abundant erect or
suberect stiff long hairs. Color blackish-
brown; hairs blackish-brown; wings rather
smoky. (Note: In the holotype, the right
antenna is abnormal, with 11 segments,
while the left one has 12.)
Comments. Pristomyrmex pollux is
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
known from Malaya and Sabah. It must
have evolved from a P. picteti—like ances-
tor. The workers of this species can be im-
mediately recognized by possessing a pair
of distinct propodeal spines that are long,
strongly upcurved at their apices (i.e.,
hooklike), and laterally compressed. This
character is unique in the genus Pristo-
myrmex.
Material Examined (MCZC, USNM,
ANIC, BMNH, NHMV). N. Borneo: Mt.
Dubit, 3,000 ft (E. Mjoberg).
Biological Information. Unknown.
Pristomyrmex reticulatus Donisthorpe
Figures 255-256
Pristomyrmex reticulatus Donisthorpe, 1949: 750.
Holotype male, New Guinea: Finschhafen,
27.iv.1944 (E. S. Ross) (CASC) [examined].
Male. TL 4.86, HL 0.82, HW 0.83, CI
HORS SIMOlS 47S Ale BOS Shee Wao 4s
AL 1.64 (n = 1).
Head, including the eyes, distinctly
broader than long. Clypeus convex in mid-
dle; its anterior margin transverse. On the
mesoscutum, notauli forming a Y shape,
with several transverse rugae. Scuto-scu-
tellar sulcus wide, separated into small
cells by several longitudinal ridges. Pro-
podeum armed ante a pair of fairly long
spines. Metapleural lobes somewhat
rounded. Petiole node in profile longer
than high with a long anterior peduncle.
Postpetiole in profile Fouled dorsally, in
dorsal view subquadrate. Dorsum of head
with a developed rugoreticulum, except
for a narrow, long median strip that is
smooth and unsculptured. Clypeus with
some longitudinal rugae. Promesonotum
with strongly developed rugoreticulum.
Sides of petiole with rugoreticulum. Sides
of postpetiole with a few longitudinal ru-
gae as well as a few superficial foveolate
punctures between them. Gaster unsculp-
tured, smooth, and shining. All dorsal sur-
faces weft numerous long hairs; hairs stiff
on the head and alitrunk. Body and. hairs
blackish-brown; wings dusky.
Comments and Discussion. This species,
described from a single male, belongs to
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
es
5295
BBS Ax LoS
ASTI <a
ae 256 we
Figures 255-256. Pristomyrmex reticulatus Donisthorpe. 255A: Male head, full-face view; 255B: Male antenna; 256: Male,
lateral view.
the umbripennis group because it is very
similar to the males of P. picteti, P. pollux,
and P. umbripennis in the structure and
shape of propodeal spines and petiole and
in body size, sculpture and hairs.
This male differs from the males of the
other three species (P. pollux, P. umbri-
pennis, and P. picteti) of the umbripennis
group as follows: In P. pollux, the head, in
full-face view, is rather narrow and long
(CI = SO) with numerous small foveolate
punctures; the mesoscutum possesses
some scattered, small and shallow foveo-
late punctures; the postpetiole in dorsal
view is distinctly longer than broad. But in
P. reticulatus, C1 is 101, the dorsal head
and the mesonotum have a developed ru-
goreticulum, and the postpetiole in dorsal
view is subquadrate.
The male of P. umbripennis is larger
(HL 0.94, HW 0.98, EL 0.47, PW 1.28,
AL 1.98, and TL 6.04) than that of P. re-
ticulatus. In addition, the dorsum of the
head between the eyes is sculptured with
a rugoreticulum in P. reticulatus but is
rather smooth in P. umbripennis.
The male of P. picteti possesses a few to
some small foveolate punctures on the
dorsal surfaces of the head and the me-
sonotum, in contrast with a rugoreticulum
in P. reticulatus.
Whether P. reticulatus represents the
male of P. fuscipennis is so far unknown.
Thus, P. reticulatus is tentatively main-
tained as a valid species until enough evi-
dence is obtained.
Ecological Information. Unknown.
Pristomyrmex umbripennis (F. Smith)
Figures 257-260
Myrmica umbripennis F. Smith, 1863: 21. Holotype
queen, Indonesia: Mysol (A. R. Wallace) (OXUM)
[examined].
Pristomyrmex umbripennis (F. Smith) Donisthorpe,
1932: 471.
Solenopsis laevis F. Smith, 1865: 75. Holotype work-
er, Indonesia: Morty Island (A. R. Wallace)
(OXUM) [examined]. Syn. n.
Pheidologeton laevis (F. Smith) Emery, 1922: 213.
Pristomyrmex laevis (F. Smith) Donisthorpe, 1932:
473.
Pristomyrmex parumpunctatus Emery, 1887: 452.
Lectotype worker, New Guinea: Andai (L. M.
D’Albertis) (MCSN), here designated, [examined].
Syn. n.
Pristomyrmex castor Donisthorpe, 1944: 81. Lecto-
type queen, New Guinea: Papua, Kokoda, 1,200 ft,
x.1933 (L. E. Cheesman) (BMNH), here designat-
ed, [examined]. Syn. n.
Pristomyrmex castaneicolor Donisthorpe, 1949: 412.
Syntype workers, New Guinea: Maffin Bay, ix-1944
(E. S. Ross) (BMNH, CASC, LACM, MCZC,
USNM) [examined]. Syn. n.
Diagnosis (Worker). Masticatory margin
of mandible with four teeth (an apical + a
preapical + two broad-based short teeth
of similar size), lacking a distinct diastema;
526
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
=
orn 4
[leap
ex
Seho/
Figures 257-260. Pristomyrmex umbripennis (F. Smith). 257A: Worker head, full-face view; 257B: Showing a transverse ridge
on the ventral clypeus; 258: Worker, lateral view; 259A: Queen head, full-face view; 259B: Showing a transverse ridge on the
ventral clypeus; 260: Queen, lateral view.
eyes generally with six to seven (rarely five)
ommatidia in the longest row; propodeum
with a pair of elongate triangular teeth.
Worker “bl > 48=.06) HE 1k32=1k63:
Ele s0S1h 74 G19 8109S 1202152:
SIRS 92. En OM =OMSS PW OrS6—1e 6s
AL 1.46=1.88, PPW 0.33-0.47, PPL 0.32-—
O44 ERE 10321139 G@r = 52):
Mandibles with a few longitudinal rugae
that often reach to the masticatory margin.
Basal margin of mandible with a central,
broadly curved prominence. Clypeus
smooth and shining with a strong median
longitudinal carina. Anterior clypeal mar-
gin usually with seven denticles: a median
denticle and three others on each side;
sometimes one to two lateral denticles are
indistinct; sometimes, two lateral denticles
are fused into a broad lobe. Ventral surface
of clypeus with a coarse transverse ridge.
Palp formula 1,3. Frontal carinae extend-
ing to the level of the posterior margins of
eyes. Antennal scrobes absent. F rontal
lobes present, partially covering the con-
dylar bulbs of holding antennal scapes.
One third to one half of the antennal
scapes usually laterally compressed near
the bases. Antennal scapes, when lying on
the dorsal head, just beyond the occipital
margin. Lamella encircling the base of the
antennal scape with a broad and deep
notch on the center of the dorsal surface.
Eyes generally containing more than 20
(rarely 15) ommatidia, with six to seven
(rarely five) in the longest row. Occipital
margin straight or feebly concave in full-
face view. Alitrunk, in profile, with a con-
vex dorsum, in dorsal view without any su-
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 527
tures. Pronotum unarmed. Propodeum
with a pair of slightly elongate triangular
teeth. Metapleural lobes prominent and
rounded. Petiole node in profile longer
than high with a long anterior peduncle;
its anterodorsal angle is on approximately
the same level as the posterodorsal. Sub-
petiole with a narrow, semitranslucent la-
mella. Postpetiole in profile higher than
long, rounded dorsally, in dorsal view
slightly broadening from front to back.
Dorsal surfaces of head and alitrunk and
the sides of pronotum with scattered fo-
veolate punctures, varying from shallow,
small, and few to distinct, rather large and
many; space between foveolae usually
smooth. Petiole, postpetiole, and gaster
smooth and shining. Dorsal surfaces of
head and alitrunk with some erect or sub-
erect short hairs. Usually two to three pairs
of hairs present on the dorsal surfaces of
petiole node and postpetiole, respectively.
A row of forward-projecting hairs present
near the anterior clypeal margin. Scapes
and tibiae with numerous suberect short
hairs. First gastral tergite lacking erect or
suberect hairs. Color reddish-brown, but
the masticatory margin of mandible black-
brown.
Queen. TL §.00=8.25, AL 1.80—2.02,
BIW 21002=222 55 C99 SS Sw o2 16a)
Si 7S Ws), 1BIL, O4O O42, JEW. IL XO EGO),
AL 2.26—2.40, PPW 0.52—0.52, PPL 0.48—
0.50, PPI 104-108 (n = 4).
General shape as in Figures 259-260,
with normal caste differences from the
conspecific worker; propodeal armaments
toothlike: other characters similar to work-
er.
Male. A single male specimen (BMNH),
collected in Papua (Kokoda, 1,200 ft) by
L. E. Cheeman in August 1933, has the
following measurements: TL ca. 6.04, HL
0.94, HW 0.98, CI 104, SL 0.35, SI 36,
EL 0.47, PW 1.28, AL 1.98.
Head, including the eyes, distinctly
broader than long. Clypeus narrow, trans-
verse, convex in the middle, its anterior
margin almost straight. Frontal carinae
short. Eyes large and prominent. On the
mesoscutum, notauli rather wide, forming
a Y shape with several coarse rugae. Scuto-
scutellar sulcus wide, separated into small
cells by longitudinal ridges. Propodeum
armed with a pair of strong triangular
teeth. Metapleural lobes rounded. Petiole
node in profile low, distinctly longer than
high, with a long anterior peduncle. Post-
petiole in profile slightly longer than high
and rounded dorsally. Dorsum of head be-
hind the level of the posterior margins of
eyes with some foveolate-reticulate sculp-
ture, but the centrical disc of the dorsal
head, under the median ocellus, smooth
and shining. Each side of the dorsal head,
between the eye and the frontal carina,
with several transverse rugae; spaces be-
tween the rugae smooth and shining. Pro-
mesonotum with somewhat coarse foveo-
late-reticulate sculpture. Sides of petiole
node with foveolate-reticulate sculpture.
Postpetiole rather smooth, only with few
superficial short rugae on each side. Gas-
ter unsculptured, smooth, and_ shining.
Body blackish-brown, but gaster and
scapes reddish-brown.
Note: This male is assigned to the spe-
cies P. umbripennis for the following rea-
sons: (1) It belongs to the wmbripennis
group because it is similar to the males of
P. picteti, P. pollux, and P. reticulatus but
different from the other known males of
Pristomyrmex in the structure and shape
of propodeal spines, petiole, notauli, and
scuto-scutellar sulcus and in body size,
sculpture, and hairs. (2) It cannot be
placed in the other species of the umbri-
pennis group. It differs from the male of
P. pollux because the former has a wider
head (HW 0.98, CI 104) than the latter
(HW 0.68—0.70, CI 80). This male is larger
(HW 0.98, HL 0.94, EL 0.47) than the
males of P. picteti and P. reticulatus (HW
< 0.85, HL < 0.85, EL < 0.40). In fact,
it is the largest male specimen so far found
in Pristomyrmex. This matches with the
workers and queens of P. umbripennis,
which are the largest in the genus. (3) The
collecting locality and the collector name
528
for this male are the same as the other two
queens of P. umbripennis.
Comments. Pristomyrmex umbripennis
occurs in New Guinea and some islands of
Indonesia. It is closely related to P. pollux
and P. picteti, but their workers and
queens can be separated by the following
characters:
P. umbripennis
Eyes larger, generally consisting of 20 or
more ommatidia and containing six to
seven (rarely five) ommatidia in the
longest row (worker)
Propodeum with a pair of triangular
toothlike armaments that are shorter
than the distance between their bases
(worker and queen)
One-third to one-half of the antennal
scape usually laterally compressed
near the base (worker and queen)
P. pollux and P. picteti
Eyes smaller, generally consisting of 10
or less ommatidia, and containing two
to three (rarely four) ommatidia in
the longest row (worker)
Propodeum with a pair of fairly long or
long spines that are longer (or much
longer) than the distance between
their bases (worker) or with a pair of
short spines (queen)
Antennal scape not laterally compressed
near the base (worker and queen)
Material Examined (ANIC, MCZC,
LAMN, BMNH, BMHB). Papua New
Guinea: Gulf Prov., Ivimka Camp, Lake-
kamu Basin, 7.7°S, 146.8°E, 120 m, #96-
266 (R. R. Snelling); NETH. Santani, 90+
m (T. C. Maa): NE Eloa River 488 to 518
m (S. Cutleck); N. Dist. of Papua, Kokoda
(P. M. Room); N. Dist. of Papua, Saiho (P.
M. Room); N. Dist. of Papua, Lejo Rd (P.
Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
M. Room); Papua, Brown Riv., lowland
rainfor., ground, rotten tree stump (R. W.
Taylor); Maffin Bay (E. S. Sepik Province,
ca. 2 to 3 km S of Wirui, S of Wewak 50
tO ICO) tm, ORBES, WaS.S7le CR. J. Ko-
hout); Lae, Bupu River, wet rotten log (B.
B. Lowery); Bulolo, 3,000 ft, rainforest, in
rotten log (B. B. Lowery); near Popondet-
ta, <50 m, rainforest, ex trunks and low
foliage (R. W. Taylor); 4 mi S of Popon-
detta, rainforest (B. B. Lowery); Sangara
N.D. (G. Baker). Indonesia: Amboina
(Bird); Irian Jaya, 50 km S Manokwari, Ar-
fak Mtns. Nature Reserve 25 m, second
rainforest, ex log (G. D. Alpert); Irian Jaya,
PT. Freeport Concession, Siewa camp.,
03.04°S 136.38°E, 200 ft, lowland second-
ary rainforest, #98-62 and #98-86, under
loose bark of log (R. R. Snelling); Seram
I., Solea (M. Brendell).
Ecological Information. This species oc-
curs in rainforest and has been collected
in and on rotten logs and under loose bark
of a log.
NOMEN NUDUM
Pristomyrmex parvispina Emery
Emery (1900) mentioned the name
Pristomyrmex parvispina but provided
neither an indication such as “n.sp.” (or its
equivalent, ele, Spe nov. 9)snor a descrip-
tion and designated no types. Pristomyr-
mex parvispind 1s thus a nomen nudum.
Emery (1922: 233) cited P parvispina
Emery as a synonym of P. brevispinosus
Emery, but this was incorrect because P.
parvispina was not a valid name. In addi-
tion, Emery (1922: 233) cited the original
publication date incorrectly as 1901 [see
Bolton’s (1995) catalog for the dating of
the original paper].
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 5
Figure 261. Head of the male of Pristomyrmex brevispinosus Emery, full-face view.
Figure 262. Head of the male of Pristomyrmex ?flatus (see the text), full-face view; mandible indicated by an arrow.
530 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
a al
we :
Figure 263. Head of the male of Pristomyrmex longispinus sp. n., full-face view.
Figure 264. Head of the male of Pristomyrmex orbiceps (Santschi), full-face view.
266
0.1 mm
Figure 265. Head of the male of Pristomyrmex quadridentatus (André), full-face view.
Figure 266. Head of the male of Pristomyrmex sulcatus Emery, full-face view.
OX
>)
1
532 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figure 267. Head of the male of Pristomyrmex obesus Mann, full-face view.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 533
Figure 268. Head of the male of Pristomyrmex picteti Emery, full-face view.
Figure 269. Head of the male of Pristomyrmex pollux Donisthorpe, full-face view (antennae missing).
53: Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figure 270. Male of Pristomyrmex brevispinosus Emery, lateral view.
Figure 271. Male of Pristomyrmex ?flatus (see the text), lateral view.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 535
1 mm
Figure 272. Male of Pristomyrmex longispinus sp. n., lateral view.
Figure 273. Male of Pristomyrmex orbiceps (Santschi), lateral view.
536 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figure 274. Male of Pristomyrmex quadridens Emery, lateral view.
Figure 275. Male of Pristomyrmex quadridentatus (André), lateral view.
Figure 276. Male of Pristomyrmex sulcatus Emery, lateral view.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang 537
<toa
RN. » en
Figure 277. Alitrunk and petiole node of the male of Pristomyrmex levigatus Emery, lateral view.
Figure 278. Head, alitrunk and petiole node of the male of Pristomyrmex obesus Mann, lateral view.
538 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figure 279. Male of Pristomyrmex picteti Emery, lateral view; propodeal spine and petiole node indicated by an arrow, re-
spectively.
Figure 280. Male of Pristomyrmex pollux Donisthorpe, lateral view; propodeal spine and petiole node indicated by an arrow,
respectively.
REVISION OF THE ANT GENUS PRISTOMYRMEX * Wang
qa
Figure 281.
539
Mesonotum of the male of Pristomyrmex levigatus Emery, dorsal view.
Figure 282. Mesonotum of the male of Pristomyrmex pollux Donisthorpe, dorsal view.
ACKNOWLEDGMENTS
I am most grateful to Dr. E. O. Wilson.
Throughout this research, I have received
constant advice, support, encouragement,
and help from him. He has taken respon-
sibility for loans of the specimens for this
research. He also reviewed two drafts of
my manuscript and provided valuable
comments.
I express my sincere thanks to Dr. James
Traniello for his valuable advice and sup-
port and for the helpful comments and
corrections he provided on an early draft
of my manuscript; to Mr. Stefan Cover for
valuable advice and discussions, for im-
proving the English expression in the text
and for other help; and to Dr. Ernst Mayr,
who has kindly answered my many ques-
tions on evolutionary biology and _syste-
matical theory. The support of Boston
University is also gratefully acknowledged.
Special thanks go to Mr. Barry Bolton.
He has provided me with valuable advice
and useful information, helped me borrow
specimens from several museums, and
sent me a printout of his catalog of Pris-
tomyrmex before his book was published.
Thanks are also due to the following
people for loans of material or other help:
Dr. Gary Alpert, Dr. James Carpenter, Dr.
Brain Farrell, Dr. Brain F isher, Ms. Kathy
Horton, Dr. Lars-Ake Janzon, Dr. F rank
Koch, Dr. Ivan L6bl, Mr. Christopher
O'Toole, Dr. Phil Perkins, Dr. Naomi
Pierce, Dr. Wojciech J. Pulawski, Dr. Al-
exander Radchenko, Dr. Valter Raineri,
Dr. Hamish G. Robertson, Dr. Louis Roth,
Dr. Ted R. Schultz, Dr. Steven O. Shat-
tuck, Dr. Stefan Schédl, Dr. David R.
Smith, and Mr. Roy R. Snelling. Finally, I
wish to thank my wife Xiufen Wang for
support and assistance.
This work was supported by E. O. Wil-
son Foundation and E. O. Wilson Evolu-
tionary Biology Fund.
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