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Full text of "Bulletin of the Museum of Comparative Zoology at Harvard College"

iS 



HARVARD UNIVERSITY 
-^ 

Library of the 

Museum of 

Comparative Zoology 



(US ISSN 0027-4100) 



The Systematics of Neotropical 

Orb-weaving Spiders in the 

Genus Metepeira (Araneae: Araneidae) 



WILLIAM H. PIEL 



( 



MCZ 
LIBRARY 

JUL 3 2001 



HARVARD 
UNIVERSITY 



HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 157, NUMBER 1 
8 JUNE 2001 



(US ISSN 0027-4100) 



PUBLICATIONS ISSUED 

OR DISTRIBUTED BY THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

HARVARD UNIVERSITY 



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SPEGIAL 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 Thoreaus 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. 

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THE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS IN 
THE GENUS METEPEIRA (ARANEAE: ARANEIDAE) 



WILLIAM H. PIEU 

CONTENTS 



Abstract _ ___. 1 

Introduction 2 

Acknowledgments 2 

Materials and Methods 3 

Collections Examined 3 

Locality Data Storage and Manipulation 4 

Examination and Illustration 4 

Metepeira F. O. P.-Cambridge 5 

Key to Female Metepeira 12 

Key to Male Metepeira 17 

Metepeira foxi Group 19 

1. Metepeira datona ChamberHn and Ivie 20 

2. Metepeira desenderi Baert 21 

3. Metepeira grandiosa grandiosa 
Chamberlin and Ivie 23 

4. Metepeira grandiosa alpina 
Chamberlin and Ivie 24 

Metepeira vigilax Group 26 

5. Metepeira cajahamba New Species .... 26 

6. Metepeira glornerabilis (Keyserling) ... 28 

7. Metepeira vigilax (Keyserling) 30 

8. Metepeira rectangula (Nicolet) 32 

Metepeira labyrinthea Group 33 

9. Metepeira spinipes F. O. P.-Cambridge .. 34 

10. Metepeira lacandon New Species 37 

Metepeira nigriventris Group 38 

11. Metepeira nigriventris (Taczanowsld) 38 

12. Metepeira tarapaca New Species 40 

13. Metepeira calamuchita New Species .. 42 

14. Metepeira galatheae (Thorell) 43 

15. Metepeira karkii (Tullgren) 46 

Metepeira conipsa Group 47 

16. Metepeira compsa (Chamberlin) 48 

17. Metepeira roraima New Species 53 

18. Metepeira gressa (Keyserling) 54 

Metepeira incrassata Group 56 

19. Metepeira maija New Species 56 

20. Metepeira inca New Species 58 



' Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts 02138. Current 
address: Institute of Evolutionaiy and Ecological Sci- 
ences, Leiden University, 2311 GP Leiden, The 
Netherlands; piel@rulsfb.leidenuniv.nl. 



21. Metepeira gosoga Chamberlin and Ivie 
59 

22. Metepeira ohnec New Species 60 

23. Metepeira comanche Levi 62 

24. Metepeira pimungan New Species 62 

25. Metepeira triangidaris (Franganillo) .. 63 

26. Metepeira arizonica Chamberlin and 

Ivie 66 

27. Metepeira atascadero New Species .... 67 

28. Metepeira incrassata F. O. P.- 
Cambridge 68 

Metepeira ventura Group 71 

29. Metepeira ventura Chamberlin and 

Ivie 71 

30. Metepeira revillagigedo New Species 73 

31. Metepeira celestun New Species 74 

32. Metepeira uncata F. O. P.-Cambridge ... 76 

33. Metepeira crassipes Chamberlin and 

Ivie 77 

34. Metepeira chilapae Chamberlin and 

Ivie 78 

Metepeira ininima Group 80 

35. Metepeira petatlan New Species 80 

36. Metepeira minima Gertsch 82 

37. Metepeira pacifica New Species 84 

38. Metepeira jamaicensis Archer 86 

Literature Cited 88 

Index 91 

Abstract. Of the 39 species and three subspecies 
of the orb-weaver genus Metepeira in the Americas, 
36 species and two subspecies are known to occur 
outside of the U.S. and Canada. Yet, despite their 
conspicuous webs, diurnal foraging, and relatively 
common presence, the taxonomy of Metepeira is 
poorly understood, probably because the genitalia are 
small and difficult to distinguish. In fact, many names 
for species south of the U.S. were, at some time, in- 
correctly synonymized with the name Metepeira la- 
byrinthea. In this paper, 14 new species are named 
(Metepeira atascadero, M. cajahamba, M. calamuchi- 
ta, M. celestun, M. inca, M. lacandon, M. maya, M. 
ohnec, M. pacifica, M. petatlan, M. pimungan, M. re- 
villagigedo, M. roraima, M. tarapaca); 11 new junior 



Bull. Mus. Comp. ZooL, 157(1): 1-92, June, 2001 1 



2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



synonyms are reported (M. acostai, M. bani, M. dom- 
inicana, M. grinnelli, M. latigyna, M. perezi, M. san- 
ta, M. salei, M. seditiosa, M. vaurieorum, M. virgi- 
nensis); five cases of erroneously synonymized names 
are reversed; 22 species and two subspecies are re- 
described (M. arizonica, M. triangidaris, M. chilapae, 
M. Comanche, M. compsa, M. crassipes, M. datona, 
M. desenderi, M. galatheae, M. glo7nerabilis, M. go- 
soga, M. grandiosa alpina, M. grandiosa grandiosa, 
M. gressa, M. incrassata, M. jamaicensis, M. karkii, 
M. minima, M. nigriventris, M. rectangula, M. spi- 
nipes, M. uncata, M. ventura, M. vigilax); and a key 
to all Metepeira species is presented. In addition, sev- 
eral ecological and life history observations are re- 
ported for various species. 

INTRODUCTION 

The absence of a comprehensive revi- 
sion of Neotropical Metepeira has left the 
taxonomy of this group in shambles. Over 
the years, a fair number of species have 
been named, particularly by A. F. Archer, 
R. V. Chamberlin, and W. Ivie. However, 
these efforts have been sporadic and, for 
the most part, scant. For example, the de- 
scription of Metepeira dominicana (Ar- 
cher, 1965) provides little information oth- 
er than "form typical of Metepeira in all 
respects," a few measurements, and two 
unrecognizable figures. Even when species 
are properly described they have far less 
taxonomic value when published alone, in 
the absence of a full comparative revision. 

The poor understanding of Metepeira 
taxonomy has persisted despite great eco- 
logical and behavioral interest in this ge- 
nus. Indeed, many species are obligate or 
facultative social species and offer excel- 
lent models for investigating genetic and 
environmental factors that influence colo- 
ny formation (e.g., Uetz and Cangialosi, 
1986; Uetz et al, 1987). The monumental 
work carried out over many years by G. W. 
Uetz has made great strides in our under- 
standing of gregarious social behavior in 
spiders and in risk-sensitive foraging the- 
ory in general (e.g., Uetz, 1996). Still, in 
the absence of solid taxonomic literature, 
behavioral ecologists have been forced to 
apply informal names to their study ani- 
mals (e.g., Metepeira "atascadero" in Uetz 
[1989] or Metepeira "Species A" in Viera 



[1989]), but this practice can lead to trou- 
ble. In one case, the behavior of several 
different species was initially studied un- 
der the false assumption that they all be- 
longed to the same species (e.g., Uetz et 
al., 1982). Clearly, a strong taxonomic 
foundation is important for further biolog- 
ical work. 

Ultimately, the relatively small, indis- 
tinct genitalia and the relatively homoge- 
neous abdominal patterns are to blame for 
the weakness in our knowledge of Mete- 
peira taxonomy. Many of these species are 
undoubtedly hard to distinguish, and this 
fact has surely intimidated arachnologists 
from taking on the painful task of revising 
the group. In the absence of good distin- 
guishing characteristics, the catalogs of 
Bonnet (1957) and Roewer (1942) synon- 
ymized the names of many Neotropical 
species with the name Metepeira lahyrin- 
thea. Levi's (1977) revision of Nearctic 
species observes that M. labyrinthea is ac- 
tually limited to the eastern United States. 
One task in this revision consists of reas- 
serting the names of species that were im- 
properly synonymized and clarifying the 
diagnostic characters that are needed to 
identify them. 

ACKNOWLEDGMENTS i 

This paper is part of my Ph.D. thesis for 
the Department of Organismic and Evo- 
lutionary Biology, Harvard University. I ain 
indebted to many people for their help, 
assistance, and encouragement in this pro- 
ject. I am especially thankful for the ded- 
ication and support of my advisors, Her- 
bert W. Levi and Edward O. Wilson. I am 
grateful that my colleagues in the Depart- 
ment of Invertebrate Zoology provided 
such a pleasant place to work: Edward 
Cutler, Ardis Johnston, Laura Leibensper- 
ger, Damhnait McHugh, Diana Sherry, 
Van Wallach, and Dee Woessner, among 
others. 

Field collecting and new specimen ac- 
quisitions were made possible with the 
help of Gita Bodner, Fundacion Capacitar, 
Tim Coonan (CINP), Fred Coyle, Dawn 



Metepeira • Piel 



Fitzpatrick, Germania Jacome, Antdnia 
Monteiro, Tila Perez, George Putnam, 
Linda Rayor, Grace Smith (NAWF), and 
George Uetz. I am particularly indebted to 
George Uetz for his assistance and corre- 
spondence. 

I am thankful for the comments by 
those who read this paper — especially to 
the members on my thesis committee: H. 
W. Levi, N. E. Pierce, and E. O. Wilson. 
I am also indebted to Kathy Horton for 
her help in formatting and preparing the 
manuscript and to the Colles Fund for de- 
fraying the costs of publication. Curators 
at various institutions who lent me speci- 
mens are listed in the Materials and Meth- 
ods section. I cannot overstress the value 
of museum collections and expert curators, 
without which research in taxonomy would 
not be possible. Museum collections are 
the most important tools available for un- 
derstanding biodiversity. 

MATERIALS AND METHODS 

Collections Examined. The taxonomic 
revision was carried out on specimens bor- 
rowed from the following collections. The 
abbreviations correspond to those listed 
with each record after eveiy species de- 
scription. I am grateful to the museums, 
curators, and staff that graciously loaned 
the material. 

ADC A. Dean, Texas A&M University, 

College Station, Texas, United 
States 

AMNH American Museum of Natural 
History, New York, United 
States; N. Platnick, L. Sorkin 

BMNH Natural History Museum, Lon- 
don, England; P. Hillyard 

CAS California Academy of Sciences, 

San Francisco, California, Unit- 
ed States; C. Griswold 

CV Carlos Valderrama A.; Bogota, 

Colombia 

FSCA Florida State Collection of Ar- 
thropods, Gainesville, Florida, 
United States; G. B. Edwards 

IRSNB Institut Royal des Sciences Na- 



turelles de Belgique, Brussels, 
Belgium; L. Baert 

JAK J. A. Kochalka, Ciudad Univer- 

sitaria, Paraguay 

JEC J. Carico, Lynchburg, Virginia, 

United States 

JMM J. Maes, Leon, Nicaragua 

MACN Museo Argentino de Ciencias 
Naturales, Buenos Aires, Argen- 
tina; E. A. Maury, C. L. Scioscia 

MCN Museu de Ciencias Naturais, 
Fundagao Zoobotanica do Rio 
Grande do Sul, Porto Alegre, 
Rio Grande do Sul, Brazil; E. H. 
Buckup, M. A. L. Marques 

MCZ Museum of Coniparative Zool- 

ogy, Harvard University, Cam- 
bridge, Massachusetts, United 
States; H. W. Levi 

MECN Museo Ecuatoriano de Ciencias 
Naturales, Quito, Ecuador; Ger- 
mania Estevez Jacome 

MEG M. E. Galiano, Buenos Aires, 
Argentina 

MLJC Maria Luisa Jimenez, Centro de 
Investigaciones Bioldgicas del 
Noroeste, La Paz, Mexico 

MLP Museo de Universidad Nacional, 

La Plata, Argentina; R. F. Arro- 
zpide, C. Sutton 

MNRJ Museu Nacional, Rio de Janeiro, 
Brazil; A. Timotheo da Costa 

MNSD Museo Nacional de Historia 
Natural, Santo Domingo, Re- 
publica Dominicana; Felix Del 
Monte 

MUSM Museo de Historia Natural, 
Universidad Nacional Mayor de 
San Marcos, Lima, Peru; D. Silva 

MZSP Museu de Zoologia, Universida- 
de de Sao Paulo, Sao Paulo, SP, 
Brazil; P. Vanzolini, J. L. Leme 

MZUF Museo Zoologico de "La Spe- 
cola" Universita di Firenze, 
Florence, Italy; S. Whitman 

NRMS Naturhistoriska Riksmuseet, 
Stockholm, Sweden; T. Krones- 
tedt 

PAN Polska Akademia Nauk, Warsza- 



4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



wa, Poland; J. Proszynski, A. Slo- 
jewska, W. B. Jedryczkowski 

REL R. E. Leech, Edmonton, Alber- 

ta, Canada 

SMF Forschungsinstitut Sencken- 

berg, Frankfurt am Main, Ger- 
many; M. Grasshoff 

SR Susan Riechert, Knoxville, Ten- 

nessee, United States 

USNM National Museum of Natural 
History, Smithsonian Institution, 
Washington, D.C., United 
States; J. Coddington, S. F. 
Larcher 

ZMB Zoologisches Museum der 

Humboldt Universitat, Berlin, 
Germany; M. Moritz 

ZMUC Zoologisk Museum, Gopenha- 
gen, Denmark; H. Enghoff, N. 
Scharff 

ZSM Zoologische Staatssammlung, 

Munich, Germany 

Locality Data Storage and Manipula- 
tion. Locality data from each collection vial 
were entered into a database designed us- 
ing Glaris FileMaker Pro®. Geographic 
coordinates were added to locality data 
that lacked them using maps, USBGN gaz- 
etteers, and on-line databases (http://164. 
214.2.59/gns/html/ and http://mapping. 
usgs.gov/www/gnis/). Occasionally locality 
information was illegible or unknown or 
one of several homonymous sites. In such 
cases a reasonable, educated guess was 
inade and a "[?]" designation was append- 
ed to the locality. In some cases the itin- 
erary of a collector was reconstructed from 
other known records, and the ambiguous 
locality was assigned a coordinate halfway 
between the previous and following known 
collection sites. The locality database 
worked in concert with the mapping pro- 
gram Atlas Pro® to generate thematic 
maps on the fly. These maps helped in the 
process of delimiting species and discov- 
ering cryptic species. 

Elevation (in meters) was estimated for 
each locality that lacked this information. 
In some cases, elevation was estimated us- 



ing contour maps, such as DMAAG ONG 
aeronautical maps; in most cases, elevation 
was estimated using NOAA data with an 
on-line database server (http://phylogeny. 
harvard.edu/~piel/find.htiril). 

The enhanced locality database was 
used to reveal ecological and life history 
traits. Seasonality of species was expressed 
by plotting a circular histogram showing 
the relative amount of collecting activity 
per 5-day interval (Figs. 300-337). While 
locality dates alone cannot control for the 
seasonal activity of human collectors, these 
data at least provide an estimate of spider 
seasonal abundance, if only approximate. 
Some syiupatric species show incongruous 
seasonal abundance, which is at least some 
evidence that seasonality of spider collec- 
tors does not unduly overshadow the sea- 
sonality of the spiders themselves. 

Examination and Illustration. Speci- 
mens were examined under 80% ethanol 
in a dish with light and dark sand grains 
for specimen support. Digital photographs 
of preserved specimens were taken 
through a Nikon SMZ-10 photomicro- 
scope using a Panasonic WV-CL320 GGD 
video camera, chosen for its high sensitiv- 
ity to light. Video images were captured 
using a Quicklmage®24 digitizer and ed- 
ited on a Quadra 700 Macintosh® com- 
puter. The computer allows relatively in- 
expensive pictures to be printed rapidly on 
a 1,200 dpi Xante® Accel-a- Writer 8200 
laser printer. Digital pictures were used to 
help sort out individuals to species, to cre- 
ate publishable pictures of gross dorsal 
and ventral markings, and to aid in the il- 
lustration of genitalia. As an aid in illustra- 
tion, the digital pictures functioned as a 
camera lucida because they assured accu- 
racy when drawing the proportions of gen- 
ital parts and scle rites. Usually a digital 
picture was laid over carbon paper and an 
outline of the genitalia was transferred to 
coquille board underneath. The illustra- 
tion continued on the coquille board using 
a Staedtler OmniGhrom® pencil and a 
drafting pen with India ink and then was 
scanned at 600 dpi on a LaGie Silverscan- 



Metepeira • Piel 



ner II®. The resulting digital image was 
edited in Adobe Photoshop® and reduced 
in size to 1,200 dpi. The edited figures 
were finally arranged on plates using Can- 
vas®. 

External genital structures were manip- 
ulated with pins to reveal hidden parts. 
The terminal division on the male palp is 
hinged, so it had to be pried open to see 
the embolus and embolic apophyses prop- 
erly. In females, mating plugs had to be 
removed from epigynal openings using 
pins. Sometimes the entire epigynum was 
partly cut from the body so as to see it 
from a posterior view. 

Internal genital structures were studied 
by clearing them in clove oil and examin- 
ing them using an Olympus BH-2 com- 
pound microscope. Sketches were made 
directly on the computer in Canvas® by 
aiming the camera lucida at the computer 
monitor. While internal genital structures 
helped in the process of delimiting spe- 
cies, they did not prove to be as useful as 
external genital structures in describing 
species; thus, these working sketches are 
not figured herein. 

Measurements of the spiders were tak- 
en using a Leitz stereo dissecting micro- 
scope with a calibrated reticule. Sizes of 
leg articles, eyes, and carapace, were per- 
formed on one specimen of each sex, for 
each species. The respective localities of 
the candidate specimens were indicated in 
the descriptions. This study placed little 
reliance on spider leg measurements be- 
cause they are not usually very useful in 
spider taxonomy, and because Metepeira 
species are notorious for their variability in 
size (Levi, 1977; Piel, 1996). 

All eye sizes were reported as a ratio of 
the posterior median eye diameters to the 
diameter of every other eye type. For ex- 
ample, in the case of "ratio of eye diame- 
ters: posterior medians and anterior me- 
dians 2.0, anterior laterals 0.5, posterior 
laterals 1.0," the reader should interpret 
the anterior medians to be half the size of 
the posterior medians, and the anterior lat- 
erals to be twice the size of the posterior 



medians. Eye separations were expressed 
in terms of their own diameters, or in 
terms of the anterior lateral eyes when be- 
tween eyes of different types. Oval eyes 
were measured as an average of the lon- 
gest and shortest lengths. 

In parallel with the last revision of Me- 
tepeira (Levi, 1977), leg measurements 
were made on each article distal to the tro- 
chanter for the first leg and on the com- 
bined lengths of the patellae and tibiae for 
all remaining legs. Variation in total body 
size was provided as an average, minimum, 
and maximum of the total lengths from a 
number of mature specimens, usually cho- 
sen from a wide geographic spread. 

Metepeira F. O. P. -Cambridge 

Metepeira F. O. P.-Cambridge, 1903: 457. Type spe- 
cies by original designation M. spinipes F. O. P.- 
Cambridge 1903. The name is feminine. 

Diagnostic Abstract. Web combines bar- 
rier or scaffolding structure surrounding a 
classic araneid orb with a retreat suspend- 
ed in air (Fig. 1). Like a raccoon with its 
facial colors reversed, the eye region is 
lighter than any other part of the carapace 
(Fig. 2). The venter has a wide median 
white line set on a black background that, 
with only some exception, extends anteri- 
orly on the sternum (Fig. 3). With one ex- 
ception, the total lengths of distal leg ar- 
ticles (metatarsus and tarsus) exceed that 
of the middle articles (patella and tibia). 
The median apophysis has two distinctive 
flagella (F in Fig. 5) and, in some species, 
an easily recognizable keel (K in Fig. 5). 
The dorsal abdominal markings (the foli- 
um) look like an inverted fleur-de-lis, al- 
lowing easy recognition of the genus in the 
field (Fig. 2). 

Description and Diagnosis. For field 
ecologists, the most obvious and distinctive 
feature of Metepeira is the combination of 
orb and barrier web (Fig. 1). The barrier 
\^eh forms scaffolding around an almost 
vertical orb and supports the spiders re- 
treat, which is thus suspended away from 
any substrate. 

In contrast to most araneids, the cara- 



6 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



pace of Metepeira is lightest in the eye re- 
gion. However, this distinctive feature 
varies within the genus: in the case of M. 
rectangula (Nicolet, 1849), the hghter re- 
gion takes up ahiiost half the carapace 
(Fig. 65); in the case of Metepeira F. O. 
P.-Cambridge, 1903, the lighter region is 
usually limited to the anterior edges of the 
carapace (Fig. 2). White, downy hairs of- 
ten cover the carapace but are especially 
white and conspicuous on the lighter parts 
of the carapace outside the eye region. In 
some species, such as M. spinipes, these 
hairs make the carapace look gray or sil- 
very when the spider is alive, but dark 
brown when the spider is in ethanol. 

The eyes of Metepeira are not particu- 
larly unusual. Eye separations relative to 
eye diameters increase with spider size: 
larger spiders tend to have relatively great- 
er eye separations. In either sex, the pos- 
terior median eyes are between 1.1 and 
1.7 times the size of the anterior medians, 
and the separation between posterior me- 
dian eyes is between 0.4 and 0.7 of that 
between anterior median eyes. The sepa- 
ration between the anterior median eyes 
and the anterior lateral eyes is between 1 
and 3.7 times the size of anterior median 
eyes in males and between one and five 
times the size in females. The diameter of 
the anterior median eyes exceeds the 
height of the clypeus. 

The shape of the female abdomen rang- 
es from wider than long and rhomboid (M. 
datona. Fig. 12) to roundish (e.g., M. de- 
senderi. Fig. 20; M. rectangula. Fig. 65), 
to longer than wide and oval (e.g., M. inca. 
Fig. 169). The dorsal folium has a recog- 
nizable white fleur-de-lis pattern, usually 
on a dark background, its edges shaped by 
a wavy, zig-zag white outline (Fig. 2). The 
dorsum of live spiders is often more red- 
dish — a pigment that rapidly dissolves in 
alcohol. 

Somewhat less common among other 
araneids is the median white line on the 
venter of the abdomen (Fig. 3), which is 
present (though shortened) even in the 
most darkly pignriented species. However, 



unique among araneid genera is the com- 
bination of median white line on the ven- 
ter and median white line on a black or 
brown sternuin. Some Metepeira species 
lack a complete white line on the sternum, 
but even those, such as M. datona, that 
usually have an entirely black sternum 
nonetheless sho\v hints of white markings 
in some specimens. Characteristics found 
in the carapace, abdomen, and sternum of 
Metepeira are also found in Araneus 
koepckeorurn Levi, but this last species 
lacks the white line on the venter. 

With the exception of M. datona, and in 
some cases, M. desenderi, all Metepeira 
species have a combined metatarsus and 
tarsus that is longer than the combined pa- 
tella and tibia. This feature is unusual 
among araneids and is not found in Kaira 
O. P. -Cambridge or other likely relatives 
to Metepeira (Levi, 1977; Piel and Nutt, 
1997). 

In most species the leg articles are 
ringed, usually with brownish black on the 
distal and dorsal surfaces of each article, 
except for the patellae and tarsi which are 
usually entirely dark. In mainly tropical 
and high-altitude species, the coxae are 
mostly black (e.g.. Fig. 75), but in desert/ 
mesquite species they appear yellowish 
white (e.g.. Fig. 28). I 

Unlike many other araneids — and per- 
haps because of the small male size — the 
coxa on leg I of male Metepeira lacks the 
hook and corresponding groove typically 
found on femur II. In addition, males lack 
a tooth on the lateral side of the endite, 
and they lack a basal tooth on the palpal 
femur. The phylogenetic analysis of 
Scharff and Coddington (1997) incorrectly 
codes Metepeira as having a tooth on the 
endite. However, had the authors coded 
this character as absent, they would have 
decreased the length of their preferred 
tree because the nearest relatives hypoth- 
esized for Metepeira {Kaira, Zijgiella, and 
Singa) also lack this tooth. 

Macrosetae usually concentrate on arti- 
cles that contact other spiders during mat- 
ing or grappling. In contrast to most gen- 



Metepeira • Piel 7 



Barrier Web 



White Dorsal Light Eye Region 

Fleur-de-Lis / ^^ Median White 

Pattern x AW* ^Ma Line on Sternum 




Orb Web 



Median White 
Line on Black 
Venter 



vigilax nigriventris incrassata minima 

foxi labyrinthea compsa vcntura 




Figure 1. Web of immature Metepeira grandiosa alpina from Chihuahua, Mexico. 

Figure 2. Dorsum of adult female Metepeira crassipes. 

Figure 3. Venter of adult female Metepeira tarapaca new species. 

Figure 4. Hypothetical phylogenetic relationships among Metepeira species groups. Shaded branches indicate species groups 

that live in South America; open branches indicate species groups that live in North America, Central America, and the Caribbean. 

Abbreviations: DEA, distal embolic apophysis; K, keel of median apophysis; TA, terminal apophysis; (+), character state gain; 

(-), character state loss. 

Figures 5, 6. Male palpus. 5, mesal view, Metepeira compsa. 6, ventral view of distal embolic division, Metepeira labyrintliea 

(Hentz). 

Abbreviations: BEA, basal embolic apophysis; C, conductor; DEA, distal embolic apophysis; E, embolus; F, flagellum on median 

apophysis; K, keel of median apophysis; MA, median apophysis; TA, terminal apophysis; TO, terminal division. 

Figures 7-13. Metepeira datona Chamberlin and Ivie (sp. 1; 17°53'N, 76°19'W). 7, male palpus, mesal. 8, epigynum, posterior. 

9, epigynum, ventral. 10, male, dorsal. 11, male, ventral. 12, female, dorsal. 13, female, ventral. 

Scale bar: dorsum and venter figures 1 .0 mm. 



8 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



era related to Araneus, Metepeira has con- 
centrated macrosetae on femur I instead 
of tibia II (Scharff and Coddington, 1997). 
Female Metepeira have between two and 
five macrosetae on die anterior side of the 
femur, and between zero and seven on the 
anteroventral side. Males typically have 
inore setae than their conspecific females: 
four to nine on the anterior side and two 
to nine on the anteroventral side. Variation 
in the number of macrosetae appears to 
correlate with body size. In most species, 
the inale palpal tibia and patella each have 
two strong macrosetae (Levi, 1977, fig. 8). 

Compared with other araneid genera, 
Metepeira have rather small and similar 
genitalia, which on the one hand makes 
the genus easy to recognize, but on the 
other hand makes species tough to iden- 
tify. The small epigynum is fleshy, variable 
in shape, and weakly sclerotized. Unlike 
Araneus, Metepeiras scape never has a 
pocket but always ends with a pointed tip 
(e.g.. Fig. 31). The cleared epigynum — 
and in many cases the uncleared epigyn- 
um — reveals a pair of sclerotized spherical 
structures where the embolus is inserted, 
as well as ducts to pass semen to the larger, 
spherical seminal receptacles. In some 
species, these spherical structures are wide 
apart (e.g.. Figs. 16, 17), in others they are 
tubular (e.g.. Figs. 39, 40), but in many, 
they are closer together (e.g.. Figs. 93, 94). 
Frequently the deeper, large seminal re- 
ceptacles can be seen through uncleared 
tissue (e.g.. Figs. 201, 295). 

The male palp is more distinctive. In 
particular, the median apophysis (MA in 
Fig. 5), while not always a good character 
for separating closely related species, is ex- 
cellent when it comes to identifying the 
genus. Two flagellae (F in Fig. 5) grace- 
fully curve off the base of the median 
apophysis, and in some species, a toothed 
or smooth keel (K in Fig. 5) extends in the 
opposite direction. This design is also seen 
in Kaira, Aculepeira, and Amazonepeira, 
but none of these have flagellae that ap- 
pear so integral to the base structure. 

The tenninal division on the Metepeira 



palp is very similar in almost all species. 
When this structure is pulled up, a basal 
embolic apophysis (also known as an em- 
bolar lamella) can be seen in the shape of 
a club or spatula (E in Figs. 5, 6). Soiue- 
times a distal embolic apophysis can be 
seen if it is not hidden from view by an 
overhanging terminal apophysis. When the 
terminal apophysis is large and sclero- 
tized — which is die case in all but the Me- 
tepeira foxi species group — it has a rec- 
ognizable toothed notch, like the mouth 
on a wrench (Fig. 6). Virgin males have a 
cap on the embolus that remains in the 
epigynum after mating and presumably 
serves as a barrier to subsequent mating 
(Levi, 1977). The shape of the embolus 
cap varies from tiny (e.g.. Fig. 178) to 
short but wide (e.g., Fig. 199) to large and 
winged (Fig. 46). Finally, the terminal di- 
vision lacks a stipes — a sclerite between 
the radix and the embolus that is frequent- 
ly found in other genera related to Ara- 
neus (Scharff and Coddington, 1997). 

Natural History. All Metepeira species 
build a unique web that combines an orb 
with a barrier web (Levi, 1977; Lopez, 
1993). As with Cijrtophora Simon or Me- 
cynogea Simon (Levi, 1997), the retreat of 
Metepeira hangs in the air, away from sub- 
strate, and is suspended by a scaffolding 
structure created by the barrier web (Fig. 
1). The spider detects vibrations in the 
web and gains quick access to the hub us- 
ing a signal line that runs from the retreat 
to the center of the orb (Fig. 1). Tan col- 
ored egg sacs are strung together, usually 
above the retreat, and the most recently 
laid eggs are nearest to the spider. In some 
species the egg sacs and retreat are deco- 
rated with insect parts (e.g., M. spinipes); 
in other species they are carefully wrapped 
by leaves and woven together (e.g., M. da- 
tona). Unlike the webs oi Cijrtophora and 
Mecynogea, the orb web of Metepeira is 
oriented vertically, and the number of radii 
and sticky spirals are more typical of other 
araneines. 

In some species, such as M. pimungan 
(personal observation) and, to a lesser de- 



Metepeira • Piel 



gree, M. incrassata (G. Uetz, personal 
communication), juveniles and adults 
without eggs will live on webs lacking a 
suspended retreat. Instead, the spider sits 
on a white disk-shaped stabilimentum in 
the center of the hub. Of 110 M. pimun- 
gan specimens observed on San Miguel Is- 
land, about 40% occupied webs of this 
type. In two cases the disk stabilimenta 
were partly separated from the hub by 
barrier web lines and were further bent 
over to form a partly covered protective 
retreat for the spider. This observation 
makes it possible to imagine that the disk 
stabilimentum seen in M. pimiingan re- 
sults from the fusion of the suspended re- 
treat with the hub. 

When food supplies are plentiful, spi- 
ders of all kinds show an increased toler- 
ance for one another and an increased ten- 
dency to aggregate (e.g., Gillespie, 1987; 
Rypstra, 1986). The suspended retreats 
and barrier webs of Metepeira, Cyrtopho- 
ra, and Mecynogea may actually further fa- 
cilitate in the formation of aggregations by 
easing dependency on substrate availabili- 
ty and by providing a common support sys- 
tem (Burgess and Witt, 1976; Uetz, 1986). 
In any case, colony formation is known to 
occur in all three genera (e.g., Rypstra, 
1979), but especially in Metepeira. Small 
colonial aggregations of two to 10 individ- 
uals occur in M. datona (Spiller and 
Schoener, 1989), M. minima (personal ob- 
servation), M. glomerabilis (R. Baptista, 
personal communication), and M. atascad- 
ero new species (e.g., Uetz and Hodge, 
1990). Medium-size colonies of 10 to 30 
individuals occur in M. pimungan (person- 
al observation), M. gressa (Viera and Cos- 
ta, 1988), M. nigriventris (L. Rayor, per- 
sonal coinmunication), M. tarapaca (V. 
Roth, locality label), and M. spinipes (e.g., 
Uetz, 1988a). Large colonies, sometimes 
in the thousands of individuals, commonly 
occur in M. incrassata (e.g., Uetz and 
Hodge, 1990). Near rivers and in other 
lush habitats, M. tarapaca colonies can 
reach 200 individuals (M. Roy, personal 
communication). These cases of social be- 



havior, broadly spread across seven differ- 
ent species groups, may mean that aggre- 
gation is a frequently lost and relatively old 
trait, or it may mean that species are prone 
to converge and evolve the same behavior 
independently. 

Either way, much research has focused 
on elucidating the selective forces behind 
colonial behavior in Metepeira. In partic- 
ular, Uetz (1988a,b, 1996) has provided 
strong support for the hypothesis that Me- 
tepeira forage using a risk-sensitive strat- 
egy. He suggests that spiders in abundant 
habitats seek to minimize individual vari- 
ance in prey capture by aggregating in col- 
onies, whereas spiders in poor habitats 
seek to maximize variance by living soli- 
tarily — perhaps in a risky attempt to find 
areas of local prey maxima. The diversity 
of social tendencies among species is 
therefore commensurate of the diversity of 
ecological habitats that they inhabit. 

Indeed, Metepeira species thrive in a 
wide array of habitats, though often they 
are quite harsh. These include wet, mon- 
tane cloud forests in Mexico and Panama 
(M. incrassata, M. olmec); tropical and wet 
agricultural areas (M. uncata, M. vigilax, 
M. glomerabilis, M. roraima); high-eleva- 
tion pine forests (M. lacandon, M. nigri- 
ventris, M. grandiosa alpina); Canadian 
bogs (M. grandiosa palustris); deciduous 
forests in the eastern U.S. (M. labyrin- 
thea); Caribbean coastal shrubbery (M. da- 
tona, M. minima, M. triangularis, M. ja- 
maicensis, M. maya, M. celestun); Mexican 
mesquite grasslands (M. atascadero, M. 
chilapae); Patagonian dunes and scrub (M. 
galatheae) and pampas grass (M. karkii); 
diy Californian buckwheat and sage (M. 
crassipes, M. ventura, M.foxi, M. grandio- 
sa grandiosa); and arid and semiarid de- 
serts (M. arizonica, M. inca, M. ventura, 
M. crassipes). Although some species (e.g., 
M. galatheae, M. spinipes, M. cornpsa) cov- 
er vast geographic areas and live in many 
different habitats, many species are more 
biogeographically restricted. In fact, sev- 
eral species follow narrow ecological zones 
that decrease in elevation with distance 



10 Bulletin Museum of Comparative Zoology, Vol. 157, No. 



from the equator (e.g., M. rectangula, M. 
vigilax, M. cajahamha. Fig. 36; M. arizon- 
ica. Fig. 213). 

Close cohabitation with different inter- 
and intrageneric species is not uncommon. 
Colonies of M. incrassata are known to 
contact webs of Nephila clavipes Linnaeus 
(Hodge and Uetz, 1996) and Mecijnogea 
ocosingo and Gasteracantha cancriformis 
(personal observation). Often M. crassipes, 
M. Ventura, M. foxi, and M. grandiosa 
grandiosa are collected together (Levi, 
1977), as are M. minima and M. celesiun 
(personal observation). Species that have 
been collected from identical localities, 
though not necessarily at the same time, 
include: M. chilapae and M. spinipes; M. 
chilapae and M. atascadero; M. karkii and 
M. galatheae; M. calamuchita new species, 
M. gressa, and M. galatheae; M. rectan- 
gula, M. calamuchita, and M. galatheae; 
M. compsa and M. gressa; M. vigilax and 
M. compsa; M. glomerahilis and M. vigilax; 
M. compsa and M. glotnerabilis; M. comp- 
sa and M. nigriventris; M. compsa and M. 
mcfl; M. datona and M. jamaicensis; and 
M. datona and M. triangularis. 

Despite the wide biogeographic ranges 
of M. compsa (Puerto Rico and south to 
Argentina, Map 8) and M. datona (His- 
paniola and north to Florida, Map 1), they 
nonetheless come geographically close to 
one another but do not overlap. It is hard 
to imagine that the hurricanes that fre- 
quently pass through the Caribbean, as 
well as the homogeneous island environ- 
ments, would not gradually cause these 
two species distributions to overlap. Per- 
haps these abrupt, disjunct distributions 
are a rare example of competitive exclu- 
sion in Metepeira, which in other species 
is not thought to be an important factor 
(Wise, 1983). 

Sphecid wasps are predators on Mete- 
peira. Locality labels indicated that M. pa- 
cifica has been found in the nests of Try- 
pargilum nitidum, T. tenoctitlan, and T. 
hensoni. Jimenez and Tejas (1994) report 
that M. crassipes is the most frequent prey 
item in the nests of Trypargilum triden- 



tatum. Colonial spiders, such as M. incras- 
sata, are especially vulnerable to wasps, 
other spiders, sarcophagid flies (e.g., Ar- 
achnidomyia Undue, A. rayorae), and 
hummingbirds (Hieber and Uetz, 1990; 
Lopez, 1989; Rayor and Uetz, 1990). 

Species Groups. Nearctic Metepeira 
were divided into two species groups: the 
M. labyrinthea group and the M. foxi 
group, based on the pattern on the ster- 
num and the shape of the median apoph- 
ysis (Levi, 1977). Baert (1987) questioned 
the taxonomic usefulness of the M. foxi 
species group (M. foxi, M. grandiosa, M. 
datona) because he found that M. desen- 
deri has both a keel on the median apoph- 
ysis and a white sternal line (Figs. 15, 
21) — a combination that is incoinpatible 
by Levi's scheme. Nonetheless, the geni- 
talia of M. desenderi closely ally this spe- 
cies with the M. foxi group, so I am re- 
defining the M. foxi group based on purely 
genitalic characters. This is likely to be a 
basal, paraphyletic group (Fig. 4) (Piel and 
Nutt, 1997). 

Seven additional species groups are dis- 
tal to the M. foxi. These remaining species 
are united by sharing a large terminal 
apophysis that is sclerotized and usually 
studded with teeth or denticles. The M. 
vigilax group (M. vigilax, M. cajahamha, 
M. glomerahilis, M. rectangida) are united 
by large emboli with long scooplike basal 
embolic apophyses (Fig. 60). Unlike the 
remaining species, the terminal apophysis 
in this group — albeit large — does not ac- 
tually overhang or hide the embolus. In 
addition to an overhanging terminal 
apophysis, the remaining taxa are also 
united by a distal embolic apophysis that 
either protrudes (Fig. 76), curves off (Fig. 
185), or is secondarily lost (Fig. 264). The 
M. labyrinthea group (M. labyrinthea, M. 
lacandon, M. spinipes) share a toothless, 
smooth keel on the median apophysis 
(Figs. 67, 69). 

The M. nigriventris group and the M. 
compsa group together share a median 
apophysis with teeth on the face of the 
keel (Figs. 92, 149). The M. incrassata 



Metepeira • Piel 



11 




Map 1 




• glomerabilis 
+ cajabamba 



Map 3 



Map 2 




grandiosa alpina 
grandiosa grandiosa' 



4»^ 




• vigilax 
+ rectangula 

Map 4 



Maps 1, 2. Metepeira fox; species group. 1, M. datona, M. desenderi. 2, M. grandiosa grandiosa. M. grandiosa alpina. 
Maps 3, 4. Metepeira vigilax species group. 3, M. glomerabilis. M. cajabamba. 4, M. vigilax, M. rectangula. 



group, the M. ventura group, and the M. 
minima group all lack a keel on the me- 
dian apophysis (Figs. 164, 222, 293). How- 
ever, both the M. compsa group and the 
M. incrassata group have epigyna with 
similar oval or round sclerotized rims 
(Figs. 151, 166), so it is likely that these 
are paraphyletic and consist of species 
leading up to a major North American 
(without a keel) and South American (with 
a keel) phylogenetic split (Fig. 4). 

The South American branch includes 
the M. compsa group (M. compsa, M. ro- 
raima, M. gressa) and, more distally, the 
M. nigriventris group (M. nigriventris, M. 
tarapaca, M. calamuchita, M. galatheae, 
M. karkii). This latter group is united by a 



distinctive and derived scape, which pro- 
jects out and down, creating a noticeable 
arch and overhang (Fig. 86). 

The remaining species all lack a keel on 
the median apophysis, and with one ex- 
ception {M. inca), they live exclusively in 
North America (from the Caribbean and 
Panama to Nevada). The M. incrassata 
group (M. gosoga, M. maya, M. inca, M. 
Comanche, M. olmec, M. atascadero, M. ar- 
izonica, M. incrassata, M. triangularis, M. 
pimungan) are very likely paraphyletic. 
The epigynum on each species seems au- 
tapomorphic and difficult to unite with any 
others. Some species (M. gosoga, M. maya, 
M. inca) have a pointed or projecting distal 
embolic apophysis (Fig. 171). Others have 



12 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



a distal embolic apophysis that curves off 
sharply but does not project forward (Fig. 
185). Finally, others have a distal embolic 
apophysis that curves off gently, almost to 
the point of hiding the existence of an 
apophysis (Fig. 206). 

The M. minima group (M. jamaicensis, 
M. minima, M. pacifica, M. petatlan) and 
the M. Ventura group (M. uncata, M. Ven- 
tura, M. celestun, M. chilapae, M. revilla- 
gigedo new species, M. crassipes) are unit- 
ed by derived characters: both have thin 
flagellae arising from a thin base on the 
median apophysis (Fig. 264), and both 
share the secondary loss of the distal em- 
bolic apophysis. The M. minima group is 
clearly monophyletic; its species all have 
their flagellae further set off from the me- 
dian apophysis on a separate, narrow stalk 
(Fig. 286). 

Key to Female Metepeira 

1 Epigynal openings strongly sclerotized, 
round but tilted so that they appear 
oval from a ventral view (Figs. 40, 48, 
55) 2 

— Epigynal openings weakly sclerotized 

(e.g.. Fig. 208), and if round, they are 
not tilted and do not appear oval from 

a ventral view (Fig. 131) 4 

2(1) Epigynal openings shaped like the en- 
trance to a snail shell (Fig. 55); His- 

paniola, Brazil, Argentina (Map 4) 

( 7 ) vigilax 

— Epigynal openings tubular (Figs. 40, 48) 

3 

3(2) Epigynal openings strongly tilted (Fig. 

40); Peru (Map 3) (5) cajabamba 

— Epigynal openings weakly tilted (Fig. 48); 

Colombia to Brazil (Map 3) 

(6) glomerabilis 

4(1) Epigynal openings large and gaping, cre- 
ating large atria inside (Figs. 61, 62); 

Chile and Argentina (Map 4) 

( 8 ) rectangula 

— Epigynal openings not gaping and not 

creating large atria (e.g.. Fig. 69) 5 

5(4) Weak posterior lobes on the epigynum 
create a single, wide epigynal depres- 
sion (Figs. 9, 17, 24, 31) or epigynum 
with crescent-shaped sclerotized open- 
ings on either side of a thin scape 
(Levi, 1977, fig. 87). Dark, sclerotized 
spheres below epigynal openings are 
greatly separated (Figs. 8, 16) 6 

— Stronger posterior lobes on the epigynum 



create separate, smaller epigynal de- 
pressions that are not crescent-shaped, 
or if crescent-shaped, the scape is thick 
and puffy (e.g.. Figs. 86, 102, 143, 166, 
187, 208). Dark, sclerotized spheres 
below epigynal openings are closer to- 
gether (e.g.. Figs. 101. 142) 11 

6(5) Sternum with longitudinal white line 
(Fig. 21); Galapagos Islands (Map 1) 
(2) desenderi 

- Sternum entirely black or brown (Figs. 

13, 28, 35) 7 

7(6) Abdomen wider than long (Fig. 12); Flor- 
ida to Hispaniola (Map 1) (1) datona 

- Abdomen longer dian wide (Figs. 27, 34) 

8 

8(7) Coxae as black as sternum (Levi, 1977, 
fig. 98); Canada-U.S. border (Levi, 

1977, map 2) 

(Levi, 1977: 212) grandiosa palustris 

— Coxae yellow or orange and lighter than 

sternum (Figs. 28, 35) 9 

9(8) Epigynum with crescent-shaped sclero- 
tized openings on eitlier side of die 
scape (Levi, 1977, fig. 87); western 
U.S. and Canada (Levi, 1977, map 2) 
(Levi, 1977: 210) foxi 

— Epigynum with wide, transverse depres- 

sion (Figs. 23, 24, 30, 31) 10 

10(9) Scape wide and stubby (Fig. 24); Baja 
California north to Canada (Map 2) 
(3) grandiosa grandiosa 

- Scape triangular (Fig. 31); in mountains 

from north-central Mexico to Canada 

(Map 2) (4) grandiosa alpina 

11(5) Scape diickness equal to greater than 
width of epigynal depressions (e.g.. 
Figs. 86, 119, 131, 143, 180, 201, 266) ' 
12 

— Scape narrower than epigynal depres- 

sions, or epigynal depressions in the 
shape of longitudinal slits (e.g.. Figs. 

238, 252, 280) 35 

12(11) Base of scape originates anteriorly and 
projects ventrally before cui'ving pos- 
teriorly. This projection creates an 
overhang and a noticeable gap between 
the scape and the genital openings 
(e.g.. Figs. 85, 86, 101, 102, 119, 143) 
13 

— Scape does not create a noticeable gap or 

overhang (e.g.. Figs. 77, 78, 130, 131, 

165, 166, 265) 18 

13(12) Rim of epigynal depressions slightly 
sclerotized and oval-shaped (Fig. 143); 
northern Brazil, French Guiana, and 
Colombia (Map 8) (17) roraima 

- Epigynal depressions without distinct rim 

(Fig. 123) or not sclerotized (e.g.. Fig. 

102) 14 

14(13) Sternum black, coxae mostly black (Fig. 



Metepeira • Piel 



13 



A calamuchita 



karkii 



mgnventris f ( ^ 



Map 5 




▲ galatheae 
• tarapaca 



Map 7 





• spinipes 
■ lacandon 



Map 6 




+ compsa 
• roraima 
■ gressa 



Map 8 



Maps 5, 7. Metepeira nighventris species group. M. calamuchita, M. I<arl<ii, M. nigriventris, M. galattieae, M. tarapaca. 
Map 6. Metepeira labyrintfiea species group. M. spinipes, M. lacandon. 
Map 8. Metepeira compsa species group. M. compsa, M. roraima, M. gressa. 



91), and carapace without lighter me- 
dian mark (Fig. 90); high altitudes in 
Bolivia and Peru near Lake Titicaca 

(Map 5) (11) nigriventri.s 

Sternum with median white line (Fig. 
99), or if sternum is black, then either 
the carapace has a median lighter ar- 
row-shaped mark (Fig. 113) or the cox- 
ae are mostly yellow (Figs. 114, 128) 
15 



15(14) Stenium brown to black with parallel 
lines on either side of median white 
line on venter: the parallel lines are 
tliicker anteriorly than posteriorly (Fig. 
128). Lower lip on epigynum thick and 
bulbous (Fig. 123); southern Argentina 
and southern Chile (Map 5) .... (15) karkii 

— Sternum with median white line, or if 

sternum is entirely brown to black, 
then parallel lines on either side of 



14 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



white line on venter are either absent 
(Fig. 114) or equally thick anteriorly as 
posteriorly (Fig. 99). Epigynum with- 
out thickened lower lip (Figs. 94, 102, 

118, 119, 120) 16 

16(15) Dark, sclerotized spheres in epigynal 
openings small (Figs. 119, 120). Ante- 
rior lip of epigynum rounded off, 
sometimes with openings shifted pos- 
teriorly and wiinkled portion of the 
scapes hood shifted more to the epi- 
gynum proper (Fig. 118); Chile and 
Argentina (Map 7) (14) galatheae 

— Dark, sclerotized spheres in epigynal 

openings large, and anterior lip of epi- 
gynum not rounded off. Openings al- 
ways located midway down the epigyn- 
um (Figs. 94, 102) .._._ 17 

17(16) General shape of epigynum is triangular 
with posterior width greater than an- 
terior width. Scooped-out depressions 
project more posteriorly than laterally 
(Fig. 102); north-central Argentina 
(Map 5) (13) calamiichita 

— General shape of epigynum square. 

Scooped-out depressions project more 
laterally than posteriorly (Fig. 94); 
northern Chile and southern Peru 

(Map 7) (12) tarapaca 

18(12) Epigynal openings almond-shaped, not 
noticeably sclerotized, and at a 40° to 
60° angle from axis of spider (Figs. 69, 
78) 19 

— Epigynal openings not almond-shaped 

(e.g.. Figs. 180, 187, 201), or if al- 
mond-shaped, slightly sclerotized and 
at an angle from spider's axis of 80° to 

100° (Figs. 151, 159, 166) 22 

19(18) Almond-shaped openings created by de- 
pression where scape arises from epi- 
gynum (Fig. 173); California, Arizona, 

northwestern Mexico (Map 9) 

(21) gosoga 

— Almond-shaped openings created by 

membranes inside depressions and not 
associated with scape (Figs. 69, 78) ..... 20 
20(19) Distinct C-shaped depression created 
where die scape arises from the epi- 
gynum (Fig. 78). Black marks inside al- 
mond-shaped openings not cross-eyed 
in appearance (Fig. 78). Black ster- 
num, usually with white spot in center 
(Fig. 83); mountainous regions of Chia- 
pas, Mexico (Map 6) (10) lacandon 

— Indistinct depression created where the 

scape arises from the epigynum (Fig. 
69). Black marks inside almond-shaped 
openings cross-eyed in appearance 
(Fig. 69). Sternum black, with or with- 
out a median white line. If with only a 
portion of a median white line present. 



usually only at the posterior end of the 
sternum (Fig. 75). Never with only one 

white spot in center 21 

21(20) Because of interspecific variability and 
polymorphism, females of the follow- 
ing two species are almost impossible 
to separate reliably without molecular 
sequence data. Small ribosomal sub- 
unit (12S) mtDNA sequence data has 
tlie following diagnostic markers. Base 
14261: ACGGT; base 14285: ATTTT; 
base 14361: ACTAC; base 14394: 
CTTAT; base 14412: ATTA. (Base 
numbers refer to homologous sites in 
tlie mitochondrion of Drosophila yak- 
uba, as reported by Clary and Wolsten- 
holme [1985].) One quarter of scape 
extends below lower lips of epigynum 
(Levi, 1977, fig. 14); New England to 
Florida and west to eastern Texas 
(Levi, 1977, map 1, but not including 

points appearing in Mexico) 

(Levi, 1977: 196) labyrinthea 

— For 12S mtDNA sequence data, the fol- 

lowing sequences are diagnostic. Base 
14261: AGGAT; base 14285: ATCTT; 
base 14361: ACCAC; base 14394: 
CTAAT; base 14412: TTTA. One third 
of scape extends below lower lips of 
epigynum (Levi, 1977, fig. 21; Fig. 69); 
Mexico City north to California (Map 

6) (9) .spinipes 

22(18) Epigynal openings small, round, and 
sclerotized (Fig. 131). Openings some- 
times hidden by wide scape (Fig. 134); 
Puerto Rico to Argentina and Chile 
(Map 8) (16) compsa 

— Epigynal openings not small, round, and 

sclerotized 23 

23(22) Rim of epigynal openings sclerotized and 
in an oval or teardrop shape (Figs. 151, 
159, 166, 187) 24 

— Rim of epigynal openings not sclerotized 

in an oval or teardrop shape (Figs. 180, 

194, 201, 208) 27 

24(23) Epigynal openings oval, small, and partly 
hidden by scape (Fig. 151); northern 
Argentina, Uruguay, and southern Bra- 
zil (Map 8) (18) gressa 

— Epigynal openings teardrop-shaped (Fig. 

187) or large and oval but not hidden 

by scape (Figs. 159, 166) _. 25 

25(24) Epigynal openings teardrop-shaped (Fig. 
187); northeastern Mexico to Texas 

(Levi, 1977, map 1; Map 11) 

(23) Comanche 

— Epigynal openings oval-shaped (Figs. 

159, 166) 26 

26(25) Lower lip of epigynum pointed (Fig. 
166), abdomen white (Fig. 169); north- 
ern tip of Peru (Map 11) (20) inca 



Metepeira • Piel 



15 




+ arizonica -x 

• gosoga 

♦ incrassata 
■ pimungan 



Map 9 




pacifica 
petatlan 



Map 10 




+ Comanche 
X inca 
♦ olmec 



Map 11 




• uncata 

♦ crassipes 
■ revillagigedo 



Map 12 



Map 13 




■ Ventura 
▲ celestun 
• chilapae 




jamaicensis 
minima b; s 



jcmaya 
+ atascadero 
• triangularis 
Map 14 




^;:^5y-.^ 



Map 15 



Maps 9, 11, 14. Metepeira incrassata species group. 9, M. arizonica, M. gosoga, M. incrassata, M. pimungan. ^^, M. comanche, 

M. inca, M. olmec. 14, M. maya, M. atascadero, M. triangularis. 

Maps 10, 15. Metepeira minima species group. 10, M. pacifica, M. petatlan. 15, M. minima, M. jamaicensis. 

Maps 12, 13. Metepeira ventura species group. 12, M. uncata, M. crassipes, M. revillagigedo. 13, M. ventura, M. celestun, M. 

chilapae. 



— Lower lip of epigynum thickened but not 

pointed (Fig. 159), abdomen dark (Fig. 
162); southern Mexico and Belize to 

Costa Rica (Map 14) (19) maya 

27(23) Rim of epigynal openings sclerotized and 
shaped hke a pair of sunglasses (Fig. 
201); eastern Cuba and Hispaniola 
(Map 14) (25) triangularis 

- Rim of epigynal openings not sclerotized 



or, if sclerotized, nol in shape of sun- 
glasses (e.g.. Figs. 208, 224) 28 

28(27) Posterior epigynal lobes converge behind 
the scape so that epigynal depressions 
appear closed off froin ventral view 
(Figs. 180, 208, 216, 224) 29 

- Posterior epigynal lobes end before they 

disappear behind the scape so that epi- 
gynal depressions appear open poste- 



16 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



riorly from ventral view (Figs. 194, 

245, 259, 266) 32 

29(28) Scape relatively long and thin, epigynal 
depressions large and round, large 
black spheres take up almost all the 
space in the depressions (Fig. 180); 
southern Veracruz to Panama (Map 11) 
(22) olmec 

— Scape relatively short, fat, and fleshy. 

Epigynal depressions not perfectly 
round; black spheres do not take up 
most of the space in the depression 

(Figs. 208, 216, 224) 30 

30(29) Epigynum puffy, scape so thick that de- 
pressions on either side of scape ap- 
pear crescent-shaped (Fig. 208); south- 
western U.S. to central Mexico (Map 
9) (26) arizonica 

— Epigynum not puffy, scape not so fat that 

depressions become crescent-shaped 

(Figs. 216, 224) 31 

31(30) Very social. Epigynal depressions large, 
disk-shaped, with shiny-smooth scler- 
otized inner surfaces and thin posterior 
lips (Fig. 224). Sternum mostly black, 
sometimes with anterior white marks. 
Venter without U-shaped mark circum- 
scribing median white line posteriorly. 
Coxae mostly dark brown (Fig. 228); 
mainly in southern Veracruz (Map 9) 
(28) incrassata 

— Mostly solitary. Epigynal depressions 

small, oval, with reduced shiny-smooth 
sclerotized surfaces and thick posterior 
lips (Fig. 216). Sternum with median 
white line or only white mark at pos- 
terior end. Venter with U-shaped mark 
circumscribing median white line pos- 
teriorly. Coxae mostly yellow (Fig. 
221); central Mexican plateau (Map 

14) (27) atascadero 

32(28) Small square-shaped epigynal depres- 
sions on either side of scape (Figs. 259, 
266) 33 

— Large, more rounded epigyniil depres- 

sions (Fig. 194) or with straight to S- 
shaped edges mostly covered by scape 

(Fig. 245) 34 

33(32) Dark spheres inside epigynal depressions 
appear slightly walleyed. Scape rela- 
tively narrower at base: about the 
width of the depressions (Fig. 266); 
southeast and south-central Mexico 
(Map 13) (34) chilapae 

— Dark spheres inside epigynal depressions 

appear slightly cross-eyed. Scape rela- 
tively wider at base: about twice the 
width of the depressions (Fig. 259); 
northwestern Mexico and California 

(Map 12) (33) crassipes 

34(32) Large, rounded epigynal depressions 



with large black spheres inside (Fig. 
194). Sternum black with posterior 
dewdrop-shaped mark. Fair of short 
parallel lines on either side of ventral 
median white line (Fig. 198). Carapace 
with large anterior white region (Fig. 
197); San Nicolas Island off southern 
California (Map 9) (24) pimungan 

— Epigynal depressions with straight to 

slightly S -shaped edges, mostly hidden 
by a wide triangular scape (Fig. 245). 
Sternum with wide median white line. 
No parallel white lines on either side 
of ventral median white line (Fig. 249). 
Carapace with small anterior white re- 
gion (Fig. 248); Yucatan Peninsula 

(Map 13) (31) celestun 

35(11) Epigynal depressions wider than long 

(Figs. 231, 252) 36 

— Epigynal depressions longer than wide 

(e.g.. Figs. 238, 273, 280, 288) 37 

36(35) Black comma shapes inside epigynal de- 
pressions (Fig. 231). Sternum with me- 
dian white line (Fig. 235). Dorsum 
lightly pigmented (Fig. 234); north- 
western Mexico and coastal California 
(Map 13) (29) Ventura 

— Black S-shaped marks inside epigynal de- 

pressions (Fig. 252). Sternum black 
with dewdrop mark at posterior end 
(Fig. 256). Dorsum darkly pigmented 
(Fig. 255); Guatemala and Costa Rica 

(Map 12) ___ (32) uncata 

37(35) Epigynal depressions indistinct anterior- 
ly. Black comma-shaped marks inside 
depressions and covered by translucent 
membranes (Fig. 238); Isla Socorro of 
the Archipielago de Revillagigedo 
(Map 12) (30) revillogigedo 

— Epigynal depressions distinct anteriorly. 

Black spheres shifted laterally and lo- 
cated outside the depressions (e.g.. 

Figs. 273, 280, 295) 38 

38(37) Epigynal depressions slit-shaped and 
usually narrower than scape (Figs. 280, 
281); northwestern Mexico and Yuca- 
tan Peninsula (Map 15) (36) minima 

— Epigynal depressions oval and wider than 

scape (Figs. 273, 288) 39 

39(38) Dark spheres larger than epigynal open- 
ings (Fig. 273); west coastal Mexico 
(Map 10) (35) petatlan 

— Dark spheres smaller than epigynal open- 

ings (Figs. 288, 295) 40 

40(39) V-shaped ridge under die scape. Dark 
spheres located behind the junction 
where the ridge meets the lateral edge 
of the epigynal depressions (Fig. 288); 

Honduras to Costa Rica (Map 10) 

(37) pacifica 

— Straight ridge under the scape. Dark 



Metepeira • Piel 



17 



spheres located laterally and outside of 
the junction where the ridge meets the 
lateral edge of the epigynal depressions 
(Fig. 295); Cayman Islands, Jamaica, 
and Haiti (Map 15) .— (38) jamaicensis 

Key to Male Metepeira 

1 Terminal apophysis tliin, small, fleshy, 
without teeth or sclerotized parts (Figs. 
7, 15, 22, 29) 2 

- Terminal apophysis enlarged, meaty, with 

teeth or sclerotized parts (e.g.. Figs. 

38, 60, 84, 199) 7 

2(1) Terminal apophysis narrow; embolus 
curled clockwise like the tip on a cork- 
screw (Figs. 7, 15) 3 

- Terminal apophysis wide; embolus tilted 

up and L-shaped (Figs. 22, 29) 4 

3(2) Curled embolus with a ridge on the up- 
per surface and raised on a pedicel; 
basal embolic apophysis enormous 
(Fig. 15). Sternum witli wide median 
white line (Fig. 21); Galapagos Islands 
(Map 1) (2) desenderi 

- Curled embolus smooth on its upper sur- 

face and not raised on a pedicel; basal 
embolic apophysis hardly noticeable 
(Fig. 7). Sternum entirely black (Fig. 
11); Florida to Hispaniola (Map 1) - -- 

( 1 ) datona 

4(2) L-shaped embolus at an acute (< 90°) an- 
gle (Levi, 1977, figs. 91-93); western 
U.S. and Canada (Levi, 1977, map 2) 
(Levi, 1977: 210) /oxi 

- L-shaped embolus at angle of 90° or 

greater (Figs. 22, 29) 5 

5(4) Median apophysis widi rounded projec- 
tion on dorsal side; jagged posterior 
edge of keel (Figs. 22, 29) 6 

- Median apophysis without projection — 

flat on dorsal side; rounded posterior 
edge of keel (Levi, 1977, fig. 105); 
along the Canadian and U.S. border, 
north to Nova Scotia and British Co- 
lumbia, south to Maine and North Da- 
kota (Levi, 1977, map 2) 

(Levi, 1977: 212) grandiosa palustris 

6(5) Lower, transverse part of L-shaped em- 
bolus longer than vertical part (Fig. 
29); mountains from north-central 

Mexico to Canada (Map 2) 

(4) grandiosa alpina 

- Lower, transverse part of L-shape em- 

bolus shorter than or equal to vertical 
part (Fig. 22); Baja California north to 
Canada (Map 2) .. (3) grandiosa grandiosa 
7(1) Terminal apophysis does not entirely 
overhang the embolus. Embolus long 
and robust, wdth a long and thin gap 
created between the embolus and the 



basal ennbolic apophysis (Figs. 38, 46, 
53, 60) 8 

- Terminal apophysis often overhangs the 

embolus, covering it from view. Em- 
bolus not long and robust, without 
long, thin gap between embolus and 
basal embolic apophysis (e.g.. Figs. 67, 

79, 92, 121, 141, 185) 11 

8(7) Longer flagellum as thick as shorter fla- 
gellum. Keel short, slim, and feather- 
shaped. Embolus as wide as base of 
median apophysis (Fig. 53); Hispanio- 
la, Brazil, Argentina (Map 4) .... (7) vigilax 

- Longer flagellum thicker than shorter fla- 

gellum. Keel absent (Fig. 38) or wide 
and arrowhead-shaped. Embolus thin- 
ner than base of median apophysis 

(Figs. 38, 46, 60) 9 

9(8) Keel absent or greatly reduced (Fig. 38); 

Pei-u (Map 3) (5) cajabamba 

- Keel present (Figs. 46, 60) 10 

10(9) Large embolus as long as basal embolic 

apophysis (Fig. 60). Normal embolic 

cap; Chile and Argentina (Map 4) 

(8) rectangula 

- Small embolus shorter than basal embol- 

ic apophysis, often seen with winged 
embolic cap (Fig. 46); Colombia to 

Brazil (Map 3) (6) glomerabilis 

11(7) Median apophysis with keel (e.g., Figs. 
92, 141) 

- Median apophysis without keel (e.g.. 

Figs. 157, 178, 286); all North Ameri- 
can or Caribbean species except for M. 

inca 

12(11) Keel without teeth; smooth (e.g.. Figs. 

67, 76); North America 13 

- Keel with teeth on face; rough (e.g.. Figs. 

84, 92, 100); South America 15 

13(12) Distal embolic apophysis sleek, pointed, 
and feather-shaped when viewed from 
underside of terminal division (Fig. 
70); Mexico City north to California 
(Map 6) (9) spinipes 

- Distal embolic apophysis spoon-shaped 

(Fig. 6) or widened with bump (Fig. 

79) 14 

14(13) Distal embolic apophysis spoon-shaped 
(Fig. 6). Sternum reddish brown with 
median white line. New England to 
Florida and west to eastern Texas 
(Levi, 1977, map 1, excluding points in 
Mexico) (Levi, 1977: 196) labijrinthea 

- Distal embolic apophysis widened with 

bump (Fig. 79). Sternum black with or 
without faint white mark in center 
(Fig. 81); mountainous regions of Chia- 
pas, Mexico (Map 6) (10) lacandon 

15(12) Distal embolic apophysis a simple exten- 
sion that projects forward, parallel to 
the embolus. Keel usuallv rounded 



12 



22 



18 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



(Fig. 129); Puerto Rico to Argentina 
and Chile (Map 8) (16) compsa 

— Distal embolic apophysis raised up or 

projected away from the embolus. Keel 
usually pointed or jagged (Figs. 84, 92, 

100, 121, 141, 149) 16 

16(15) Distal embolic apophysis points up and 
away from the embolus. Keel usually 
jagged (Figs. 141, 149) 21 

— Distal embolic apophysis wide and raised 

up on boss. Keel usually pointed (Figs. 
84, 92, 100) 17 

17(16) Dewlap extension under embolus (Fig. 
100) curves under, narrowing the gap 
between the embolus and the basal 
embolic apophysis by one-half (Fig. 
103); north-central Argentina (Map 5) 

(13) calarnuchita 

No such dewlap (Figs. 84, 92, 110, 121). 
Gap between embolus and basal em- 
bolic apophysis narrow by less than 
one-half the widest distance (Figs. 87, 
95, 117, 124) 18 

18(17) Outside edge of distal embolic apophysis 
gently rounded when viewed from un- 
derside of terminal division (Fig. 124); 
southern Argentina and southern Chile 
(Map 5) (15) karkii 

— Outside edge of distal embolic apophysis 

with distinct bump when viewed from 
underside of terminal division (Figs. 

87, 95, 117) 19 

19(18) Outside edge of distal embolic apophysis 
with rounded bump that is sclerotized 
and has a distinct line rising up from 
embolus (Fig. 117); Chile and Argen- 
tina (Map 7) (14) galatheae 

— Outside edge of distal embolic apophysis 

with pointed bump, less sclerotized 
than embolus proper and without a dis- 
tinct line rising up from the embolus 

(Figs. 87, 95) 20 

20(19) Thick neck joining embolus and basal 
embolic apophysis. Bump on distal em- 
bolic apophysis peeks out from under 
the terminal apophysis (Figs. 92, 95). 
Sternum usually with white spot or me- 
dian white line (Fig. 97); northern 

Chile and southern Peru (Map 7) 

(12) tarapaca 

— Thin neck joining embolus and basal em- 

bolic apophysis. Bump on distal em- 
bolic apophysis does not peek out from 
under the terminal apophysis (Figs. 84, 
87). Sternum always black, general col- 
oration dark (Figs. 88, 89); high alti- 
tudes in Bolivia and Peru near Lake 

Titicaca (Map 5) (11) nigriventris 

21(16) Distal embolic apophysis thinner tlian 
embolus near die junction of the two. 
Embolus tip is curved gendy (Fig. 



149); northern Argentina, Uruguay, 
and southern Brazil (Map 8) ... (18) gressa 

— Distal embolic apophysis the same size as 

embolus near the junction of the two. 
Embolus tip is curved abruptly (Fig. 
141); northern Brazil, French Guiana, 

and Colombia (Map 8) (17) roraima 

22(11) Distal embolic apophysis a projecting 

bump (Figs. 157, 164, 171) 23 

— Distal embolic apophysis curved off 

(Figs. 178, 185, 199, 214), rounded 
(Figs. 192, 206), or absent (Figs. 264, 

278, 293) 25 

2.3(22) Distal embolic apophysis raised up from 
the embolus and pointed (Fig. 171); 
California, Arizona, northwestern Mex- 
ico (Map 9) (21) gosoga 

— Distal embolic apophysis not raised up, 

but projecting forward and rounded off 

(Figs. 157, 164) 24 

24(23) Main flagellum on median apophysis thin 
and initially as thick as base. Embolus 
and embolic cap shortened (Fig. 164); 
northern tip of Peru (Map 11) ... (20) inca 

— Main flagellum on median apophysis 

thick, but initially thinner than base. 
Embolus and embolic cap elongated 
(Fig. 157); southern Mexico and Belize 

to Costa Rica (Map 14) (19) mnija 

25(22) Embolus shaped like the nib on a foun- 
tain pen, with bump near tip on op- 
posite side of distal embolic apophysis 
(Figs. 214, 217); central Mexican pla- 
teau (Map 14) (27) atascadero 

— Embolus witliout such bump near tip 

(e.g.. Fig. 222) 26 

26(25) Distal embolic apophysis abruptly ends 
in sharp curve and flagellae not set off 
on a narrow stalk (Figs. 178, 185, 199, 
222) 27 

— Distal embolic apophysis gently cui-ved 

off (Figs. 229, 236, 243), rounded 
(Figs. 192, 206), or absent (e.g.. Fig. 

293) 30 

27(26) Flagellae veiy thin and equal in length. 
Embolus long and arching, with wide 
but short embolus cap. Overhanging 
terminal apophysis covers only a distal 
portion of the sclerotized part of the 
embolus (Fig. 199); eastern Cuba and 
Hispaniola (Map 14) (25) triangularis 

— Flagellae normal in thickness and usually 

of different lengths. Terminal apophy- 
sis centered above the entire sclero- 
tized portion of the embolus (Figs. 

178, 185, 222) 28 

28(27) Not known to be very social. Height of 
embolus plus distal embolic apophysis 
just before the latter curves off sharply 
is equal to or greater than length of 
embolus tip distal to this point (Figs. 



Metepeira • Piel 



19 



178, 185). Sternum with median white 
markings (Figs. 182, 189) 29 

- Highly social species. Height of embolus 

plus distal embolic apophysis just be- 
fore die latter curves off shai^ply is less 
than the length of the embolus tip dis- 
tal to this point (Fig. 222). Sternum 
mostly black; sometimes with anterior 
white marks. Coxae mostly dark brown 
(Fig. 226); mainly in southern Vera- 
cruz, Mexico (Map 9) (28) incrassata 

29(28) Darker, sclerotized portion of the embo- 
lus does not extend over the hump of 
the distal embolic apophysis. Base of 
embolus narrower than widest part of 
the first flagellum (Fig. 178); southern 
Veracruz, Mexico to Panama (Map 11) 
(22) olmec 

- Darker, sclerotized portion of the embo- 

lus extends over the hump of the distal 
embolic apophysis. Width of embolus 
base the same or greater than widest 
part of the first flagellum (Fig. 185); 
northeastern Mexico to Texas (Levi, 

1977, map 1; Map 11) (23) comanche 

30(26) Distal emboUc apophysis rounded off to 

form convex curve (Figs. 192, 206) 31 

- Distal embolic apophysis gently falls off 

to form concave shape (Figs. 229, 236, 

243) or absent (e.g.. Fig. 293) 32 

31(30) Embolus S-shaped (Fig. 192). Sternum 
black with posterior white mark. Ven- 
ter without white anchor shape inark 
(Fig. 196); San Nicolas Island off 

southern California (Map 9) 

(24) pimiingan 

- Embolus convex on upper surface, 

straight on lower surface (Fig. 206). 
Sternum black with median white line. 
Venter with faint white anchor-shaped 
mark posterior to median white line 
(Fig. 210); southwestern U.S. to cen- 
tral Mexico (Map 9) (26) arizonica 

32(30) Flagellae on median apophysis set off on 
separate, narrow, stalk (e.g.. Figs. 278, 
286) 38 

- Flagellae on median apophysis not set off 

on a separate, narrow, stalk (e.g.. Figs. 

229, 264) 33 

33(32) Larger flagellum twice as wide as smaller 
flagellum (Fig. 236); Isla Socorro of the 
Archipielago de Revillagigedo (Map 
12) (30) revillagigedo 

- Larger flagellum less than twice as thick 

as smaller flagellum (e.g.. Figs. 229, 

243) 34 

34(33) Flagellae on median apophysis relatively 

thin (Figs. 250, 264) 35 

- Flagellae on median apophysis thicker 

(Figs. 229, 243, 257) 36 

35(34) Embolus with sharp bend near the tip 



(Fig. 264). Sternum with median white 
line (Fig. 268); southeast and south- 
central Mexico (Map 13) (34) chilapae 

— Embolus with gentle bend further from 

the tip (Fig. 250). Sternum mosdy 
black with small white mark at poste- 
rior end of sternum (Fig. 254); Gua- 
temala and Costa Rica (Map 12) 

(32) uncata 

36(34) Embolus with sharp bend near the tip 
(Fig. 229); northwestern Mexico and 
coastal California (Map 13) ... (29) ventura 

— Embolus with gentle bend further from 

the tip (Figs. 243, 257) 37 

37(36) Sclerotized portion of embolus about as 
long as the longer flagellum (Fig. 243); 

Yucatan Peninsula (Map 13) 

(31) celestun 

— Sclerotized portion of embolus shorter 

than the longer flagellum (Fig. 257); 
northwestern Mexico and California 

(Map 12) (33) crassipes 

38(32) Embolus thick and tapering to a point 

(Figs. 271, 278) 39 

— Embolus thin and needlelike (Figs. 286, 

293) 40 

39(38) Longer flagellum greater than half the 
length of the cymbium (Fig. 271); west 
coastal Mexico (Map 10) (35) petatlan 

— Longer flagellum less than half the length 

of the cymbium (Fig. 278); northwest- 
ern Mexico and Yucatan Peninsula 

(Map 15) (36) minima 

40(38) Flageflae set off on long, thin stalk (Fig. 
293); Cayman Islands, Jamaica, and 
Haiti (Map 15) (38) jamaicensis 

— Flagellae set off on short, diicker stalk 

(Fig. 286); Honduras to Costa Rica 
(Map 10) (37) pacifica 

Metepeira foxi Group 

The M. foxi species group {sensu Levi, 
1977) and M. desenderi share veiy similar 
genitaha, especially among males. Conse- 
quently, I am expanding the M. foxi group 
to include M. desenderi, but with the ex- 
clusion of the black sternum as a diagnos- 
tic character. The M. foxi group {sensu 
lato) includes males with an embolus that 
lacks a distal apophysis. The embolus curls 
almost 180° clockwise around a reduced or 
fleshy terminal apophysis (Figs. 7, 15, 22, 
29). In other species groups the embolus 
is straighter and does not curl (e.g.. Fig. 
60), and the terminal apophysis is large, 
overhanging the embolus, and often scler- 
otized with teeth (e.g.. Fig. 121). Epigynal 



20 



Bulletin Museum of Comparative Zoologtj, Vol. 157, No. 1 



features uniting females in the M. foxi spe- 
cies group are harder to discern. Generally 
speaking, the epigynum has a stubby, 
shorter, often triangular scape and weaker 
posterior lobes that permit a wider view 
into the epigynal openings (compare Figs. 
9, 17, 24, 31 with Figs. 143, 252). 

1 . Metepeira datona 
Chamberlin and Ivie 
Figures 7-13, 312; Map 1 

Metepeira datona Chamberlin and Ivie, 1942: 68, fig. 
196, ? . Female holotype from Daytona Beach, 
Florida, USA, in the AMNH, examined. Levi, 
1977: 208-210, figs. 78-86, S, 9. Brignofi, 1983: 
275, 

Metepeira inenna Bryant, 1945: 378. Female holo- 
type from Cap Haitian, Haiti in the MCZ, exam- 
ined. First synonymized by Levi, 1977: 208—210. 

Description. Female from Morant 
Point, Mammee Bay, Jamaica. Carapace 
light around eyes with short lateral poste- 
rior extensions, sometimes with thin lon- 
gitudinal white line (Fig. 12). Legs ringed. 
Femur I with row of two macrosetae on 
anterior side; three light setae on anter- 
oventral side. Anterior half of dorsal foli- 
um white, posterior half black, margined 
by white; two halves separated by trans- 
verse white line. Abdomen widest in cen- 
ter (Fig. 12). Venter with a longitudinal 
white line. Pair of white spots on either 
side of spiracle (Fig. 13). Sternum usually 
entirely black, though soinetimes with thin 
anterior and posterior marks, suggestive of 
median white line (Fig. 13). Ratio of eye 
diameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.3, posterior 
laterals 1.1. Anterior median eyes separat- 
ed by 1.4 diameters, posterior median eyes 
by 0.9, anterior median eyes separated 
from anterior laterals by 1.6 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.3 their diameters. Total length 4.2 
mm. Carapace 1.7 mm long, 1.4 wide. 
First femur 2 mm, patella and tibia 2.4, 
metatarsus 1.5, tarsus 0.8. Second patella 
and tibia 1.9 inm, third 1.1, fourth 1.6. 

Male from Morant Point, Mammee Bay, 
Jamaica. Carapace yellowish brown, light 



around eyes. Median white mark anterior 
to thoracic furrow (Fig. 10). Legs same 
color as carapace, lightly ringed. Femur I 
with row of three macrosetae on anterior 
side, two on anteroventral side. Center of 
dorsum with transverse white line; poste- 
rior half darker than anterior half; thin me- 
dian black line; margin of folium white 
(Fig. 10). Venter, sternum as in female 
(Fig. 11). Ratio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.3, posterior laterals 1.2. Anterior 
median eyes separated by 1.8 diameters, 
posterior median eyes by 1.1, anterior me- 
dian eyes separated from anterior laterals 
by 1.5 diameters of anterior lateral eyes, 
lateral eyes separated by 0.5 their diame- 
ters. Total length 3 mm. Carapace 1.4 mm 
long, 1.1 wide. First femur 2.7 mm, patella 
and tibia 2.7, metatarsus 1.9, tarsus 0.8. 
Second patella and tibia 2.1 mm, third 1.0, 
fourth 1.3. 

Diagnosis. The dorsal folium and ab- 
dominal shape is distinctive in M. datona. 
Typically, the females abdomen is widest 
in the middle, forming a rhomboid in 
shape (Fig. 12), and both sexes have a 
transverse white line dividing the dorsal 
folium, with wide black markings on the 
posterior half (Figs. 10, 12). A ventral view 
of the epigynum in M. datona reveals a 
ridge under the scape that almost forms a 
straight line and which cui'ves up at the 
ends (Fig. 9) instead of a V-shape (Figs. 
17, 24, 31). The openings to the epigynum 
consist of small slits that flank a wide de- 
pression (Fig. 9), compared to larger 
openings that are relatively closer together 
(Figs. 17, 24, 31). The smooth embolus on 
M. datona is somewhat more compressed 
as it curls clockwise like the tip on a cork- 
screw, in contrast to the slightly ridged 
embolus supported on a stalk (Fig. 15) or 
the strongly ridged and tilted embolus 
(Figs. 22, 29). Like M. desenderi (Fig. 15) 
but unlike M. grandiosa grandiosa (Fig. 
22) and M. grandiosa alpina (Fig. 29), tlie 
posteroventral edge of the keel on the me- 
dian apophysis is rounded, as opposed to 
pointed (Fig. 7). Finally, M. datona has the 



Metepeira • Piel 



21 



thinnest and fleshiest terminal apophysis 
in the genus (Fig. 7). 

Variation. Average body length of 15 fe- 
males examined 3.8 mm, range 2.9 to 4.8 
mm. Average body length of seven males 
examined 2.6 mm, range 1.8 to 3 mm. 

Natural History. Mature adults have 
been collected throughout the year (Fig. 
312). According to locality labels, M. da- 
tona are found near the beach on cactus, 
palm, low scrub, mangrove, and bamboo. 
Females are known to wrap their egg sacs 
in dead leaves. 

Distribution. At sea level in Florida, Ba- 
hamas, British West Indies, and the Do- 
minican Republic (Levi, 1977, map 2; Map 
1). 

Records Examined. BAHAMAS Abaco Cays: Aba- 
co, 26°29'N, 77°5'W, 2.xii.l964 (W. B. Peck, MCZ); 
Allons Cay, 26°59'N, 77°40'W, 9.V.1953 (E. B. Hay- 
den, AMNH); Hopetown, Elbow Cay, 26°33'N, 
76°57'W, 5.V.1953 (E. Hayden, AMNH); New Plym- 
outh, Green Turtle Cay, 26°50'N, 77°23'W, 7.V.1953 
(G. Rabb, AMNH); Whale Cay 26°43'N, 77°14'W, 
12.i.l964 (W. B. Peck, CAS). Acklins Id.: Atwood's 
Harbor, 22°13'N, 74°18'W, 15.ix.l958 (A. W. Scott Jr., 
MCZ); Salina Point, 22°13'N, 74°18'W, 15.viii.l958 
(R. Robertson & A. W. Scott Jr, MCZ). Andros: Fresh 
Creek, 24°26'N, 77°57'W, 23.iv.1953 (L. Giovannoh, 
AMNH); Mangrove Cay, 24°15'N, 77°39'W, 
26. iv. 1953 (E. Hayden, AMNH); Nicolls Town, 
25°8'N, 78°0'W, 14.iii.l967 (A. M. Nadler, AMNH). 
Berry Islands: Frazier's Hog Cay, 25°24'N, 77°50'W, 
29.iv.1953 (E. Hayden, AMNH); Little Harbor Cay, 
25°34'N, 77°43'W, l.vl953 (Hayden & Giovannoh, 
AMNH). Crooked Island: North shore of Cripple 
Hill, 22°49'N, 74°16'W, 15.ix.l958 (A. W. Scott Jr, 
MCZ); NW end, Gordon (= Gun) Bluff, 22°50'N, 
74°20'W, 15.viii.l958 (R. Robinson & A. W. Scott Jr, 
MCZ). Crooked Island Group: Long Cay 22°37'N, 
74°20'W, 7.iii.l953 (Hayden & Giovannoh, AMNH). 
East Plana Cay: 22°37'N, 73°.33'W, 4.iii.l953 (E. 
Hayden, AMNH). Exurnas: Musha Cay 23°50'N, 
76°15'W, 29.xii.1985 (A. Boutard, MCZ); Warderick 
Wells Cay 24°22'N, 76°36'W, 9.i.l953 (L. Giovannoh, 
AMNH), ll.i.l953 (Hayden & Giovannoh, AMNH). 
Grand Bahamas Island: Dundee Bay, 26°30'N, 
79°15'W [?], 25.xii.1965 (L. Pinter, MCZ); near Fre 
port, pine-palmetto, 26°34'N, 78°27'W [?], 
25.vii.1965 (L. Pinter, MCZ). Long Island: Clarence 
Town, 23°6'N, 74°59'W, 10.iii.l953 (L. Giovannoh, 
AMNH); Deadman's Cay, 23°14'N, 75°14'W, 
ll.iii.l953 (E. Hayden, AMNH). New Providence Is- 
land: 7 mi W of Nassau, 25°5'N, 77°28'W, 4.i.l953 
(Hayden & Giovannoli, AMNH). North Bimini: 
25°44'N, 79°15'W, 25.i.l950 (C. M. Bogert, AMNH), 



6.vi.l950 (M. Gazier & F. Rindge, AMNH), 15.V.1951 
(W. J. Gertsch & M. Gazier, AMNH), 15.vi.l951 (M. 
Gazier, P & C. Vaurie, AMNH), 15.vii.l951 (C. & P 
Vaurie, AMNH), 13.xii.l952 (A. M. Nadler, AMNH), 
28.xi. 19.59 (A. M. Nadler, AMNH). Rmn Cay: near 
Port Nelson, 2.3°38'N, 74°50'W, 16.ih.l953 (Hayden 
& Giovannoli, AMNH). South Bimini: 25°42'N, 
79°17'W, 12.vi.l950 (M. Gazier & F Rindge, 
AMNH), 22.vi.1950 (M. Gazier & F. Rindge, 
AMNH), 15.V.1951 (W. J. Gertsch & M. Gazier, 
AMNH), 15.vii.l951 (C. & P Vaurie, AMNH), 
4.vLii.l951 (C. & P Vaurie, AMNH), 4.xii.l952 (A. M. 
Nadler, AMNH), 25.ih.1953 (A. M. Nadler, AMNH); 
Gun Cay 25°35'N, 79°20'W, 15.vi.l951 (AMNH). 
BRITISH WEST INDIES Caicos Islands: Long Cay, 
21°28'N, 71°33'W 10.ii.l953 (E. Hayden, AMNH); 
South Caicos, from webs in upper beach zone, 
21°31'N, 71°30'W, 3.iv.l973 (D. W Buden, MCZ); 
West Caicos, 21°39'N, 72°28'W, 4.U.1953 (Hayden & 
Giovannoh, AMNH), 5.ii.l953 (Hayden, Rabb, & 
Giovannoli, AMNH). Grand Cayman Island: 
19°20'N, 81°10'W, 15.ii.l960 (R. A. Levidn, MCZ). 
CUBA Oriente: Banes, 20°58'N, 75°43'W, 2.viii.l955 
(A. F. Archer, AMNH); Ensenada Nispero, Giudamar, 
19°58'N, 75°52'W, 9.xi.l945 (P Alayo, AMNH); Jur- 
agua, 19°56'N, 75°40'W, l.x.1955 (P Alayo, AMNH); 
Santa Fe [?], 20°22'N, 75°53'W (A. F Archer, 
AMNH). DOMINICAN REPUBLIC Barahona: Pla- 
ya Los Patos, 17°58'N, 71°11'W, 31.viii.l976 (J. A. 
Ottenwalder, MNSD). La Altagracia: Punta Cana, 
Isla Saona, 18°8'N, 68°34'W (Felix E. Del Monte & 
K. Guerrero, MNSD). HAITI Dept. du Nord: Cap- 
Haitien, 19°46'N, 72°13'W, 15.iii.l934 (E. Bryant, 
Utawana Exp., MCZ). JAMAICA Portland: between 
Boston and Blue Hole, 18°6'N, 76°37'W, 29.vii.1955 
(A. F Archer, AMNH). Saint Andrews: Hope Gar- 
dens, Gordontown, 18°2'N, 76°45'W, 27.vii.1955 (A. 
F Archer, AMNH). Surrey: Morant Point, Mammee 
Bay, 17°53'N, 76°19'W, 14.X.1957 (A. M. Chickering, 
MCZ); Palisadoes, 17°56'N. 76°46'W, ll.xi.l967 (A. 
M. Chickering, MCZ); Palisadoes Area, 17°56'N, 
76°46'W, l.xi.l957 (A. M. Chickering, MCZ). U.S. 
VIRGIN ISLANDS Saint Thomas: Morant Point, 
17°55'N, 76°10'W, 25.vii.1985 (G. B. Edwards, 
FSCA). 

2. Metepeira desenderi Baert 
Figures 14-21, 307; Map 1 

Epeira labyrinthea:— Banks, 1902: 60. Banks, 1924: 
97. Misidentification. 

Metepeira sp.: — Roth and Craig, 1970: 116. 

Metepeira desenderi Baert, 1^87: 145, figs. 16-21, S , 
9 . Male holotype from Isla Pinzon, Galapagos, Ec- 
uador, in the IRSNB. Platnick, 1989: 341. 

Note. Holotype not examined because the figures 
in Baert (1987) are clear and because this is the 
only Metepeira species found on the Galapagos. 

Description. Female from east slope of 
Isla Santa Cruz, Galapagos Islands, Ecua- 



22 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



dor. Carapace yellowish brown, white 
around eyes, lighter median line (Fig. 20). 
Legs yellowish white, slightly darker an- 
nulations on distal ends of articles. Femur 
I with row of four macrosetae on anterior 
side; two to four on anteroventral side. 
Dorsal fleur-de-lis pattern broken into four 
white patches with anterior pair often larg- 
er than in most species. Posterior pair 
straighter than usual, forming a cross in 
center of folium (Fig. 20). Venter olive- 
brown with median white line surrounded 
by white U-shaped marking (Fig. 21). Pair 
of white spots on either side of spiracle. 
Sternum brownish black with wide, white 
line widening anteriorly (Fig. 21). Ratio of 
eye diameters: posterior medians and an- 
terior medians 1.0, anterior laterals 1.3, 
posterior laterals 1.3. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 2.3 di- 
ameters of anterior lateral eyes, lateral eyes 
separated by 0.2 their diameters. Total 
lengdi 5.5 mm. Carapace 2.3 mm long, 1.7 
wide. First femur 2.7 mm, patella and tibia 
3, metatarsus 2.2, tarsus 0.8. Second pateUa 
and tibia 2.4 mm, third 1.4, fourth 2. 

Male from same locality as female. Car- 
apace yellowish brown with wide white 
median mark (Fig. 18). Legs ringed like 
female. Femur I with row of four macro- 
setae on anterior side; four to seven on an- 
teroventral side. Dorsum and venter as in 
female, though median white line on ster- 
num sometimes broken (Figs. 18, 19). Ra- 
tio of eye diameters: posterior medians 
and anterior medians 0.9, anterior laterals 
1.2, posterior laterals 1.1. Anterior median 
eyes separated by 2.2 diameters, posterior 
median eyes by 1.4, anterior median eyes 
separated from anterior laterals by 2.3 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.5 their diameters. To- 
tal length 3.2 mm. Carapace 1.6 mm long, 
1.3 wide. First femur 2.6 nam, patella and 
tibia 2.6, metatarsus 2.3, tarsus 0.8. Sec- 
ond patella and tibia 2.2 mm, third 1.0, 
fourth 1.6. 

Diagnosis. Within the M. foxi species 




40 80 120 160 

Collection day (since Jan 1 ) 



200 



Figure 14. The length of mature M. desendeh collected on 

specific days of the year over a period of 84 years. There is a 

trend from smaller spiders early in the season to large spiders 

later in the season. 

Scale of abscissa: -80 = October 12 of the previous year; 40 

= February 9; 160 = June 9. 

Symbols: Females (o), stippled regression line (t^ = 40%); 

males (•), solid regression line (i^ = 42%). 



group, the female abdomen shape of M. 
desenderi is closest to M. datona, though 
not quite as rhomboid (compare Fig. 20 
with Figs. 12, 27, 34). The scape on M. 
desenderi is narrower and less triangular 
than other M. foxi species, and the de- 
pression under the scape forms a distinct 
U-shaped smile that is not seen in the oth- 
ers (compare Fig. 17 with Figs. 9, 24, 31). 
Like M. datona (Fig. 7) but unUke M. 
grandiosa grandiosa (Fig. 22) or M. gran- 
diosa alpina (Fig. 29), the posteroventral 
edge of the keel on the median apophysis 
is rounded as opposed to pointed (Fig. 15). 
Similar to M. datona (Fig. 7), the embolus 
of M. desenderi is curved like a slightly 
flattened corkscrew. Unlike M. datona, the 
embolus seems to be supported by a ped- 
icel off a large basal embolic apophysis 
(Fig. 15). In contrast, a pedicel is not ev- 
ident in M. datona, and the basal embolic 
apophysis is barely visible (Fig. 7). 

Variation. Average body length of 45 fe- 
males exaiTiined 5.8 mm, range 3.8 to 8.5 
mm. Average body length of 13 males ex- 
amined 4.9 mm, range 2.9 to 6.5 mm. 

Natural History. Notes on the collection 
labels of Y. Lubin and R. Silberglied in- 
dicate that M. desenderi is active at night 



Metepeira • Piel 



23 



and that the retreat is composed of Opun- 
tia bark and croton leaves. According to 
Baert (1987), M. desenderi is found in 
large numbers in arid ecological zones on 
all islands. Although some mature speci- 
mens have been collected in November 
and August, most specimens are found be- 
tween Januaiy and June (Fig. 307), cor- 
responding to the "warm and wet" season 
in the Galapagos (van der Werff, 1978). 
The size of mature M. desenderi speci- 
mens appears to correlate with the collec- 
tion date: female spiders taken in August 
are, on average, 80% larger than those tak- 
en in November (Fig. 14). This trend may 
indicate that M. desenderi can vary the 
number of instars before maturity. Alter- 
natively, the intermolt growth may be 
greater for spiders with their antepenulti- 
mate or penultimate instars occurring dur- 
ing the warmer and wetter season. 

Distribution. Endemic to the Galapagos 
Islands (Map 1). 

Records Examined. ECUADOR Galapagos: Albe- 
marle, Tagus Cove, 0°16'S 91°22'W, 23.1.1899 
(AMNH), 8.ii.l899 (AMNH), 21.iii.l899 (AMNH), 
23.iii.1899 (AMNH); Archipielago de Galapagos, 
0°0'N, 90°30'W (Williams Exped., 1923, MCZ); Ba- 
hia Conway, Indefatigable Island, 0°33'S, 90°32'W, 
17.iii.l935 (Exline-Peck, CAS); Barrington Island, 
0°49'S, 90°4'W, l.viii.l929 (H. H. Cleaves, CAS); 
Campion, nr. Floreana (Santa Maria), 1°15'S, 
90°27'W, l.vi.l981 (Y. Lubin, MCZ); Charles Island, 
1°17'S, 90°26'W, 10.V1899 (AMNH); Indefatigable 
Island, 0°38'S, 90°23'W, 27.iv.1899 (AMNH), 
18.vi.l929 (Pinchot South Sea Exp, USNM); Isla 
Abingdon, 0°35'N, 90°44'W, 25.vi.1899 (AMNH); Isla 
Albemarle, 0°30'S, 91°4'W, 13. i. 1899 (AMNH), 
20.ii.l899 (AMNH), 20.iii.l899 (AMNH); Isla Bin- 
dloe, 0°19'N, 90°29'W, 20.vi.l899 (AMNH); Isla 
Hood, 1°23'S, 89°39'W, 18.V.1899 (AMNH), 
26.V1899 (AMNH); Isla Pinta, S Coast, 0°35'N, 
90°44'W, 25.V1964 (D. Q. Cavagnaro, CAS); Isla Pla- 
za, 0°35'S, 90°9'W, 7.iii.l970 (R. Silberglied, MCZ), 
26.xi.1973 (Y. Lubin, MCZ); Isla Santa Cruz, Acade- 
my Bay, 0°44'30"S, 90°17'30"W, 13.iii.l970 (R. Sil- 
berglied, MCZ); Isla Santa Cruz, E slope, 0°38'S, 
90°23'W, 16.ivl964 (D. Q. Cavagnaro, CAS); Isla 
Santa Cruz, Estacion Cientifica Charles Darwin, 
0°44'S, 90°18'W, 24.i.l964 (D. Q. Cavagnaro & R. O. 
Schuster, CAS), 12.ii.l964 (Cavagnero & Schuster, 
CAS), 3.xi.l973 (Y. Lubin, MCZ), 24.xi.1973 (Y Lu- 
bin, MCZ), 25.xi.1973 (Y. Lubin, MCZ); Isla Santa 
Fe, S coast, 0°50'S, 90°4'W, 30.i.l983 (Y. Lubin, 
MCZ); James, 0°16'S, 90°42'W, 22.ivl899 (AMNH); 



Narborough Island, 0°25'S, 91°30'W, 28.iii.1899 
(AMNH); Sombrero Chino, Rocas Bainbridge, SE of 
Santiago, 0°21'S, 90°34'W, 31.iii.l970 (R. Silberglied, 
MCZ); Tower Island, Darwin's Bay [?], 0°17'N, 
89°59'W (AMNH); Tower Islands, 0°20'N, 89°58'W, 
7.ivl925 (N.Y. Zoological Society, AMNH); W coast 
of Albemarle Island, 0°11'N, 91°21'W, 9.iii.l935 
(AMNH). 

3. Metepeira grandiosa grandiosa 
Chamberrm & Ivie 
Figures 22-28, 321; Map 2 

Metepeira grandiosa Chamberlin and Ivie, 1941: 17, 
figs. 24—26, 9 . Female holotype from Ben Lo- 
mond, California, USA in the AMNH, examined. 

Metepeira palomara Chamberlin and Ivie, 1942: 72, 
figs. 200-204, 9, 6. Female holotype and para- 
types from Mt. Palomar, California, in the AMNH, 
examined. First synonymized by Levi, 1977: 214. 

Metepeira grandiosa grandiosa: — Levi, 1977: 214, 
figs. 112-116, ?, (J. 

Note. Levi (1977) opted to collapse M. paliistris, 
M. grandiosa, M. alpina, M. dakota, and M. palo- 
mara into three subspecies of M. grandiosa, with 
the caveat that more data may show the three sub- 
species to be distinct species. D. Buckle (personal 
communication) claims that his own recent obser- 
vations suggest that M. grandiosa alpina and M. 
grandiosa palustHs are separate species. However, 
since the bulk of M. grandiosa specimens are out- 
side of the geographic range of this revision, and 
in the absence of molecular data, I will follow 
Levi's (1977) recommendation and leave these as 
separate subspecies. 

Description. Female from Los Angeles, 
California, USA. Carapace light around 
eyes with inedian white line extending to 
thoracic furrow (Fig. 27). Legs ringed. Fe- 
mur I with row of three to four macrosetae 
on anterior; one on anteroventral. Dorsum 
with usual folium, though lighter and more 
speckled than in most species (Fig. 27). 
Venter with a wide longitudinal white line. 
Pair of white spots on either side of spi- 
racle (Fig. 28). Sternum entirely black 
(Fig. 28). Ratio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.2, posterior laterals 1. Anterior 
median eyes separated by 1.9 diameters, 
posterior median eyes by 1.3, anterior me- 
dian eyes separated from anterior laterals 
by 3 diameters of anterior lateral eyes, lat- 
eral eyes separated by 0.4 their diameters. 
Total length 6.5 mm. Carapace 3.2 mm 



24 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



long, 2.3 wide. First femur 3.5 mm, patella 
and tibia 3.7, metatarsus 3.2, tarsus 1.1. 
Second patella and tibia 3.2 mm, third 1.9, 
fourth 2.8. 

Male from Parque Nacional Sierra San 
Pedro Martir, Baja California Norte, Mex- 
ico. Carapace as in female. Legs lightly 
ringed. Macrosetae on femur I variable — 
usually row of four macrosetae on anterior 
side, five on anteroventral side. Dorsum as 
in female (Fig. 25). Venter and sternum as 
in female (Fig. 26). Ratio of eye diameters: 
posterior medians and anterior medians 
1.1, anterior laterals 1.5, posterior laterals 
1.5. Anterior median eyes separated by 1.7 
diameters, posterior median eyes by 0.8, 
anterior median eyes separated from an- 
terior laterals by 2.1 diameters of anterior 
lateral eyes, lateral eyes separated by 0.3 
their diameters. Total length 3.6 mm. Car- 
apace 1.9 mm long, 1.5 wide. First femur 
2.9 mm, patella and tibia 2.8, metatarsus 
2.5, tarsus 0.9. Second patella and tibia 2.3 
mm, third 1.3, fourth 1.9. 

Diagnosis. The epigynal scape of M. 
grandiosa grandiosa is shorter and stub- 
bier than in other members of the M. foxi 
species group (compare Fig. 24 with Figs. 
9, 17, 31). Unlike M. datona and M. de- 
senderi (Figs. 7, 15), the corkscrew em- 
bolus is tilted up with a heavy ridge (Fig. 
22), but unlike M. grandiosa alpina (Fig. 
29), it is more graceful and not as sharply 
bent. The posteroventral edge of the keel 
on the median apophysis in M. grandiosa 
grandiosa is not as pointed (Fig. 22) as in 
M. grandiosa alpina (Fig. 29) but not 
curved as in M. datona and M. desenderi 
(Figs. 7, 15). 

Variation. Body length of females varies 
from 5.4 to 8.5 mm; males from 3.5 to 5.1 
mm (Levi, 1977). 

Natural History. Mature specimens 
have been collected from March to Sep- 
tember (Fig. 321; Levi, 1977) in yellow 
pine forests and on Eriogonum fascictda- 
tum bushes. Elevations range from sea lev- 
el to 2,000 m. 

Distribution. Coastal mountainous re- 



gions from British Columbia to Baja Cal- 
ifornia Norte (Levi, 1977, map 2; Map 2). 

Records Examined. MEXICO Baja California Nor- 
te: Parque Nacional Sierra San Pedro Martir, 
30°45'N, 115°13'W, l.vii.l977 (C. E. Griswold, CAS). 
USA Arizona: Sycamore Canyon [?] Santa Cruz Co, 
31°28'N, 110°42'W, 9.ix.l978 (G. F. Knowlton, 
MCZ). California: Los Angeles, 34°3'N, 118°15'W 
(Davidson, MCZ); Winchester, Double Butte, 
33°42'N, 117°5'W, 20.iv,1974 (W. Icenogle, MCZ). 

4. Metepeira grandiosa alpina 
Chamberlin and Ivie 
Figures 29-35, 325; Map 2 

Metepeira dakota Chamberlin and Ivie, 1942: 73, figs. 
205—207, ? , c? . Male holotype and female paratype 
from Noonan, North Dakota, USA, in the AMNH, 
examined. Name synonymized by Levi, 1977: 212— 
214. 

Metepeira alpina Chamberlin and Ivie, 1942: 74. Fe- 
male holotype and female paratypes from Fish 
Lake, Utah, USA, in tlie AMNH, examined. 

Metepeira grandiosa alpina: — Levi, 1977: 212-214, 
figs. 99, 100, 106-111, 9,6. BrignoH, 1983: 276. 

ISlote. As first revisor, Levi (1977) chose to use 
the name M. alpina because its type specimen was 
collected closer to the center of the subspecies dis- 
tribution. 

Description. Female from Charcas, San 
Luis Potosi, Mexico. Carapace light 
around eyes with lateral posterior exten- 
sions and median white line extending td 
thoracic furrow (Fig. 34). Legs lightly 
ringed. Femur I with row of four macro- 
setae on anterior side; one on anteroven- 
tral side. Dorsum with usual folium, 
though lighter than in most species (Fig. 
34). Venter with a wide longitudinal white 
line. Pair of white spots on either side of 
spiracle (Fig. 35). Sternum entirely black 
(Fig. 35). Ratio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.1, posterior laterals 1.1. Anterior 
median eyes separated by 1.7 diameters, 
posterior median eyes by 0.7, anterior me- 
dian eyes separated from anterior laterals 
by 2.8 diameters of anterior lateral eyes, 
lateral eyes separated by 1.1 their diame- 
ters. Total length 6.7 mm. Carapace 3 mm 
long, 2.4 wide. First femur 3.5 mm, patella 
and tibia 3.8, metatarsus 3.4, tarsus 1.2. 



Metepeira • Piel 25 




desenderi 

(2) 




grandiosa grandiosa 
(3) 




m^ grandiosa alpina 

V (4) 



Figures 15-21. Metepeira desenderi Baert (sp. 2; 0°38'S, 90°23'W). 15, male palpus, mesal. 16, epigynum, posterior. 17, 

epigynum, ventral. 18, male, dorsal. 19, male, ventral. 20, female, dorsal. 21, female, ventral. 

Figures 22-28. Metepeira grandiosa grandiosa Chamberlin and Ivie (sp. 3 [22, 25, 26] 30°45'N, 115°13'W; [23, 24, 27, 28] 

34°3'N, 118°15'W). 22, male palpus, mesal. 23, epigynum, posterior. 24, epigynum, ventral. 25, male, dorsal. 26, male, ventral. 

27, female, dorsal. 28, female, ventral. 

Figures 29-35. Metepeira grandiosa alpina Chamberlin and Ivie (sp. 4 [29,32,33] 25°56'N, 105°22'W; [30,31,34,35] 23°8'N, 

101°7'W). 29, male palpus, mesal. 30, epigynum, posterior. 31, epigynum, ventral. 32, male, dorsal. 33, male, ventral. 34, female, 

dorsal. 35, female, ventral. 

Scale bars: dorsum and venter figures 1 .0 mm. 



26 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



10000 



1000 



■I 100 ; 




° vigilax 
X rectangula 
• cajabamaba 



JQ I I I I ''I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I 

-45 -35 -25 -15 -5 5 15 25 

Latitude (Degrees North) 

Figure 36. The elevation of collection localities for M. vigilax, 
M. rectangula, and M. cajabamba at their corresponding lati- 
tudes. Species-specific altitudes appear to decrease with dis- 
tance from the equator. Elevations estimated from NOAA da- 
tabase of 5- by 5-minute geographic tiles. Regression line of 
M. vigilax does not include data points north of the equator. 
Symbols: M. vigilax (o), M. rectangula [X], M. cajabamba [•]. 



Second patella and tibia 3.3 mm, third 1.9, 
fourth 2.9. 

Male from Santa Maria del Oro, Duran- 
go, Mexico. Carapace yellowish brown, 
light around eyes. Median white triangular 
mark anterior to thoracic furrow (Fig. 32). 
Legs same color as carapace. Macrosetae 
on femur I variable — usually row of four 
macrosetae on anterior side, five to six on 
anteroventral side. Dorsum, venter, and 
sternum as in female (Figs. 32, 33). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.0, anterior laterals 1.4, 
posterior laterals 1.2. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 0.8, anterior median eyes 
separated from anterior laterals by 2.3 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 5.2 mm. Carapace 2.5 mm long, 
2 wide. First femur 4.5 mm, patella and 
tibia 4.6, metatarsus 4.5, tarsus 1.3. Sec- 
ond patella and tibia 3.6 mm, third 1.8, 
fourth 2.8. 

Diagnosis. The epigynal scape of M. 
grandiosa alpina has the widest base of all 
members of the M. foxi species group, and 
the lobes surrounding the central depres- 
sion are soiuewhat fatter (compare Fig. 31 
witli Figs. 9, 17, 24,). Unlike M. datona 
and M. desenderi (Figs. 7, 15), the cork- 



screw embolus is tilted up with a heavy 
ridge (Fig. 29), but unlike M. grondiosa 
grandiosa (Fig. 22), it is thicker and with 
a sharper L-shape bend instead of a more 
cui'ved L-shape. The posteroventral edge 
of the keel on the median apophysis in M. 
grandiosa alpina is pointier (Fig. 29) than 
in M. grandiosa grandiosa (Fig. 22) and 
not curved as in M. datona and M. desen- 
deri (Figs. 7, 15). 

Variation. Body length of females varies 
from 4.0 to 6.8 mm; males from 3.1 to 5.3 
mm (Levi, 1977). 

Natural History. According to Levi 
(1977), mature specimens have been col- 
lected from June to August in meadows of 
bunchgrass, browsed aspen, oak, juniper, 
and sagebrush (Fig. 325). Elevations are at 
around 2,000 m. 

Distribution. North American Rockies 
from southern Alberta and Saskatchewan 
to Central Mexico (Levi, 1977, map 2; 
Map 2). 

Records Examined. MEXICO Durango: Santa Ma- 
ria del Oro, 25°56'N, 105°22'W, 28.vii.1947 (W. J. 
Gertsch, AMNH). San Luis Potosi: Charcas, moun- 
tain side, 23°8'N, 101°7'W, 7.vii.l934 (MCZ). USA. 
Colorado: Cimarron, 38°27'N, 107°33'W, 21.vii.l959 
(H. W. & L. Levi, MCZ); Hayden Creek, Sangre de 
Cristo Mtns., 38°25'N, 105°35'W [?], Il.vii.l961 (H. 
W. & L. Levi, MCZ). Utah: SE shore of Bear Lakei 
41°59'N, 111°20'W, 3.vii.l978 (G. F. Knovi^lton, 
MCZ). 

Metepeira vigilax Group 

Spiders in the M. vigilax group (Mete- 
peira cajabamba, Metepeira glome rabilis, 
Metepeira vigilax, Metepeira rectangula) 
are characterized by large, sclerotized epi- 
gynal openings, not unlike a snails shell 
(Figs. 40, 48, 55, 62), and long, robust, 
emboli with large scooplike basal embolic 
apophyses (Figs. 38, 46, 53, 60). 

5. Metepeira cajabamba new species 
Figures 36, 38-45, 315; Map 3 

Holotype. Male from Cajabamba, Cajamarca, Peru, 
25. Lx. 1955, W Weyi-auch, in CAS. The specific 
name is a noun in apposition after the localit)'. 

Description. Female paratype from Ca- 
jabamba, Cajamarca, Pei-u. Carapace dark 



Metepeira • Piel 27 



brown, light around eyes with lateral pos- 
terior extensions (Fig. 43). Proximal two- 
thirds of femora white, remainder dark 
browii. Remaining articles lightly annulat- 
ed. Femur I with row of three macrosetae 
on anterior side; none on anteroventral 
side. Dorsal folium dark; white fleur-de-lis 
pattern reduced with thin branches (Fig. 
43). Venter of abdomen black with wide 
white median line. Pair of small white 
spots on either side of colulus (Fig. 44). 
Sometimes the posterior end of median 
line ends in anchor shape (Fig. 45). Ster- 
num black, sometimes with (Fig. 44) or 
without (Fig. 45) median white line. Ratio 
of eye diameters: posterior medians and 
anterior medians 1.1, anterior laterals 1.4, 
posterior laterals 1.2. Anterior median 
eyes separated by 1.6 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 3 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 6.4 mm. Carapace 3 mm long, 
2.3 wide. First femur 3.3 mm, patella and 
tibia 3.5, metatarsus 2.9, tarsus 0.9. Sec- 
ond patella and tibia 2.9 mm, third 1.7, 
fourth 2.7. 

Male holotype. Carapace dark brown, 
white around eyes with lateral posterior 
extensions and median white mark (Fig. 
41). Proximal third of femora white, re- 
mainder dark brown. Remaining articles 
lightly annulated. Femur I with row of 
three macrosetae on anterior side; one on 
anteroventral side. Dorsal folium dark; 
white fleur-de-lis pattern indistinct with 
thin and broken branches (Fig. 41). Venter 
of abdomen black with median white 
mark. Pair of small white spots on either 
side of spiracle and colulus (Fig. 42). Ster- 
num black, often with median white line 
(Fig. 42). Ratio of eye diameters: posterior 
medians and anterior inedians 1.0, anterior 
laterals 1.3, posterior laterals 1. Anterior 
median eyes separated by 1.5 diameters, 
posterior median eyes by 0.9, anterior me- 
dian eyes separated from anterior laterals 
by 2.3 diameters of anterior lateral eyes, 
lateral eyes separated by 0.3 their diame- 



ters. Total length 4 mm. Carapace 1.9 mm 
long, 1.5 wide. First femur 3 mm, patella 
and tibia 3, metatarsus 2.7, tarsus 0.9. Sec- 
ond patella and tibia 2.4 mm, third 1.2, 
fourth 1.8. 

Diagnosis. Female M. cajahamba and 
M. glomerabilis differ from the other spe- 
cies in the M. vigilax group (M. vigilax and 
M. rectangula) by the smaller and more 
tubelike openings to the epigynum (com- 
pare Figs. 40, 48 with Figs. 55, 62). Fe- 
male M. cajabamha differs froin M. glom- 
erabilis by having epigynal openings that 
are more oval (Fig. 40) than round (Fig. 
48) when viewed ventrally, having the epi- 
gynal openings farther apart (compare Fig. 
40 with Fig. 48), and by having the scler- 
otized tubelike openings more anteriorly 
directed (Fig. 40) than parallel to the epi- 
gynal groove (Fig. 48). Male M. cajabarnba 
and M. glomerabilis differ from other spe- 
cies in the M. vigilax group by the smaller, 
thinner, and more graceful emboli (com- 
pare Figs. 38, 46 with Figs. 53, 60). Me- 
tepeira cajahamba differs from M. glom- 
erabilis by lacking a keel on the median 
apophysis, an only slightly fatter embolus, 
and having a normal embolus cap, in con- 
trast to a winged embolus cap (Fig. 46). A 
larger portion of the prosoma is dark in M. 
cajahamba (Fig. 43) as compared to M. 
glom,erabilis (Fig. 51). 

Variation. Average body length of nine 
females examined 6.2 mm, range 5.4 to 7.5 
mm. Average body length of four males 
examined 4 mm, range 3.7 to 4.9 mm. 

Natural History. With the exception of 
two specimens, mature M. cajabarnba 
specimens have been collected in May 
through October (Fig. 315). Altitudes of 
collection localities appear to correlate 
steeply with latitude (Fig. 36). Median el- 
evation, about 500 m, with a range from 
near sea level to 3,500 m. 

Distribution. Ecuador and Peru (Map 
3). 

Records Examined. PERU Ancash: Callejon de 
Huaylas, 9°10'S, 77°45'W, 15.viii.l988 (V. and B. 
Roth, CAS). Cajamnrca: Cajabarnba, 12.427, 7°37'S, 
78°3'W, 25.ix.1955 (W. Weyrauch, CAS). La Libet-tad: 



28 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Cerro Campana, La Cumbre, 8°1'S, 79°5'W, 
10.x. 1966 (A. F. Archer, AMNH). Liiiia: 20 km E An- 
con, 11°47'S, 76°59'W, l.xi.l953 (W. Weyrauch, 
CAS); 23 mi N Pativilca, 10°42'S, 77°47'W, 15.1.1955 
(E. I. Schlinger and E. S. Ross, CAS); 5 km NW 
Chilca, 12°29'S, 76°48'W, 12.ix.l954 (E. I. Schlinger 
and E. S. Ross, CAS); between moudis of Rio Chillon 
and Ancon [?], 11°54'S, 77°7'W, 5.viii.l953 (M. 
Koepcke and Koepcke, MUSM); Cajacay [?], Rio 
Fortaleza, 10°40'S, 77°52'W, 6.iii.l956 (W. Weyrauch, 
CAS); Canta, Rio Chillon, 11°28'12"S, 76°37'23"W, 
12.V.1951 (W. Weyrauch, CAS); Cerro Caracoles, 
12°23'S, 76°45'W 15.b£.1951 (W. Weyi-auch, CAS); 
Lima, 12°3'S, 77°3'W (K. Jelski and Stolzman, PAN); 
Lomas de Iguanil [?] (Huaral), 11°29'51"S, 
77°12'12"W, 14.vi.l986 (D. Silva, MUSM). 

6. Metepeira glomerabilis (Keyserling) 
Figures 37, 46-52, 314; Map 3 

Epeira glomerabilis Keyserling, 1892: 154, fig. 113, 
9 , <S . Male and female syntypes from Taquara, Rio 
Grande do Sul and Serra Vermelha, Rio de Janeiro, 
Brazil, in BMNH, examined. Male here designated 
lectotype. 

Araneus glomerabilis: — Petrunkevitch, 1911: 294. 
Roewer, 1942: 843. 

Aranea .santa Chamberlin, 1916: 254, pi. 19, fig. 10, 
9 . Female holotype from Santa Ana, 3,000 m, Cuz- 
co, Peru, in MCZ, examined. NEW SYNONYMY. 

Metepeira santa: — Chamberlin and Ivie, 1942: 67. 
Platnick, 1993: 449. 

Metepeira glomerabilis: — Chamberlin and Ivie, 1942: 
68, fig. 181. 

Metazygia gloiTierabilis: — Levi, 1991: 179. Platnick, 
1993: 448. Erroneous transfer. 

Description. Female from Punapi, Tun- 
gurahua, Ecuador. Carapace brown; white 
arrowhead mark between eye region and 
thoracic furrow (Fig. 51). Legs dusky 
brown, hghter on ventral surfaces of fem- 
ora I and II; lighter on dorsal surfaces of 
patellae, tibiae, and metatarsi I and II. Fe- 
mur I with row of two to three macrosetae 
on anterior side; none on anteroventral 
side. Dorsal folium dark, speckled white, 
and white fleur-de-lis pattern with thin 
branches (Fig. 51). Venter black with 
short, wide, white median line ending in a 
T-shape; pair of small white spots on either 
side of spiracle (Fig. 52). Sternum black 
with median white line widening anteriorly 
(Fig. 52). Ratio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.3, posterior laterals 1.2. Anterior 
median eyes separated by 1.3 diameters, 



posterior median eyes by 0.9, anterior me- 
dian eyes separated from anterior laterals 
by 1.7 diameters of anterior lateral eyes, 
lateral eyes separated by 0.2 their diame- 
ters. Total length 5 mm. Carapace 2.4 mm 
long, 1.7 wide. First femur 2.2 mm, patella 
and tibia 2.3, metatarsus 1.9, tarsus 0.9. 
Second patella and tibia 1.9 mm, third 1.2, 
fourth 1.8. 

Male from Puiiapi, Tungurahua, Ecua- 
dor. Carapace as in female, except for a 
greater separation between white colora- 
tion around eyes and arrowhead mark 
(Fig. 49). Legs light tan, dark distally on 
femora. Femur I with row of three macro- 
setae on anterior side; four on anteroven- 
tral side. Dorsal folium and venter as in 
female (Figs. 49, 50). Sternum black with 
partly broken median white line widening 
anteriorly (Fig. 50). Ratio of eye diame- 
ters: posterior medians and anterior me- 
dians 1.1, anterior laterals 1.3, posterior 
laterals 1.2. Anterior median eyes separat- 
ed by 1.8 diameters, posterior median eyes 
by 1.2, anterior median eyes separated 
from anterior laterals by 2 diameters of an- 
terior lateral eyes, lateral eyes separated by 
0.3 their diameters. Total length 3.5 mm. 
Carapace 1.7 mm long, 1.4 wide. First fe- 
mur 2.5 mm, patella and tibia 2.4, meta- 
tarsus 2.3, tarsus 0.9. Second patella and 
tibia 1.9 mm, third 1.0, fourth 1.7. 

Diagnosis. Female M. ^omerabilis and 
M. cajabamba differ from the other spe- 
cies in the M. vigilax group (M. vigilax and 
M. rectangula) by the smaller and more 
tubelike openings to the epigynum (com- 
pare Figs. 40, 48 with Figs. 55, 62). Me- 
tepeira glomerabilis differs from M. caja- 
bamba by having epigynal openings that 
are more round (Fig. 48) than oval (Fig. 
40) when viewed ventrally, having the epi- 
gynal openings closer together (compare 
Fig. 48 with Fig. 40), and by having the 
sclerotized tubelike openings more parallel 
to the epigynal groove (Fig. 48) than an- 
teriorly directed (Fig. 40). Male M. glom- 
erabilis and M. cajabamba differ from oth- 
er species in the M. vigilax group by the 
smaller, thinner, and more graceful emboli 



Metepeiba • Piel 



29 



(compare Figs. 38, 46 with Figs. 53, 60). 
Metepeira glomerabilis differs from M. ca- 
jabaniba by having a keel on the median 
apophysis, an only slightly slimmer embo- 
lus, and a larger, winged embolus cap 
(compare Fig. 46 with Fig. 38). A larger 
portion of the prosoma is white in M. 
glo77ierabilis (Fig. 51), compared to M. ca- 
jabaniba (Fig. 43). In addition, the mar- 
gins of the folium, particularly in the male, 
are whiter than in other species in the 
group (compare Fig. 49 and Keyserling 
[1892, fig. 113b] with Figs. 41, 56, 63). 

Variation. Average body length of 28 fe- 
males examined 5.1 mm, range 3.8 to 7.3 
mm. Average body length of 10 males ex- 
amined 3.1 mm, range 2.4 to 4.3 mm. The 
base of the embolus varies from relatively 
thin (Fig. 46) to somewhat thicker, as in 
M. cajabamba (Fig. 38). 

Natural History. At first it would appear 
that this species is not seasonal — mature 
specimens have been collected throughout 
the year (Fig. 315). However on closer in- 
spection, there seems to be a seasonal shift 
with elevation: mature spiders are found 
at low altitudes (0—500 m) between March 
and October, at medium altitudes (500— 
1,500 m) between August and March, and 
at high altitudes (1,500-4,000 m) between 
December and June (Fig. 37). In coastal 
regions, R. Baptista (personal communi- 
cation) reports that this species forms 
small aggregations of two to 10 spiders. 

Distribution. Colombia, Ecuador, Peru, 
Bolivia, Paraguay, and southern Brazil 
(Map 3). Elevations range froin sea level 
to 4,000 m. 

Records Examined. BOLIVIA Chuquisaca: Mon- 
teagudo, 19°49'S, 63°59'W, 24.xii.1984 (L. E. Pena, 
AMNH). BRAZIL EspiHto Santo: Fazenda Santa 
Maria [?], Apiaca, 21°4'S, 41°25'W, 22. ix. 1985 (R. L. 
C. Baptista, MZSP). Mato Grosso: Chavantina, 
14°40'S, 52°21'W, 15.vi.l947 (J. C. Carvalho, MNRJ); 
Fazenda Cei-vo, Tres Lagoas, 20°48'S, 51°43'W, 
18. ix. 1964 (Exp. Depto. ZooL, MZSP); Utiariti, 
13°2'S, 58°17'W, 15.vlii.l961 (Lenks, MZSP). Minas 
Gerais: Lavras, 21°14'S, 45°0'W, 20.x. 1978 (W. Don 
Fronk, MCZ); Pedra Azul, 16°1'S, 41°16'W, 
15.xii.l970 (F. M. Oliveira, AMNH); Peti Forest Res. 
[?] Santa Barbara, on bushes in cerrado, 19°56'S, 
43°24'W, 28.viii.1986 (R. L. C. Baptista, MZSP). Pa- 



Peru, Ecuador, Colombia 
Brazil, Paraguay, Bolivia 




50 100 150 200 250 300 350 400 450 500 550 
Collection Day (since Jan 1) 

Figure 37. The elevation of collection localities for mature spi- 
ders of M. glomerabilis on specific days of the year between 
1939 and 1990. There appears to be a shift in seasonal mat- 
uration times that corresponds better with elevation than with 
latitude. Elevations estimated from NOAA database of 5- by 
5-minute geographic tiles. 

Scale of abscissa: 1 00 = April 9; 300 = October 27; 500 = 
May 15 of the following year. 

Symbols: Peru, Ecuador, Colombia [■]; Brazil, Paraguay, Bo- 
liva [♦]. 



rand: Praia do Leste, Paranagua, 2.5°46'S, 48°31'W, 
4.V.1967 (P. Biasi, MZSP). Rio de Janeiro: Guaratiba, 
22°58'S, 42°48'W, 28.viii.1976 (J. A. R Dutra, MZSP); 
Ilha de Santana, Macae, 22°25'S, 41°44'W, 18.x. 1986 
(R. L. C. Baptista, MZSP); Itaipu, Niteroi, 22°56'S, 
43°5'W, 20.ivl985 (R. L. C. Baptista, MZSP). Rio 
Grande do Norte: Fazenda Canaan [?], Macaiba, 
5°51'S, 35°21'W, 15.ix.l951 (M. Alvorenga, MZSP). 
Rondonia: Vila Rondonia, 10°52'S, 61°57'W, 9.ii.l961 
(Pereira & Machado, MZSP). Santa Catarina: Nova 
Teutonia, 27°3'S, 52°24'W, 12.vl949 (SMF). Sao 
Paulo: Barueri, 23°31'S, 46°53'W, 13.iii.l966 (K. Len- 
ko, MZSP); Campos do Jordao, 22°44'S, 45°35'W, 
15.iii.l945 (Wygod, MZSP); Institute Oceanografico, 
Ubatuba, 2.3°26'S, 45°4'W, 1.5.vl967 (P Montouchat, 
MZSP); km 1 Rod, Rio Santos [?], Ubatuba, 23°26'S, 
45°4'W, 12.X.1985 (R. L. C. Baptista, MZSP), 
13.X.1985 (R. L. C. Baptista, MZSP); km 3 Rod, Rio 
Santos [?], Ubatuba, 23°26'S, 45°4'W, 6.i.l985 (R. L. 
C. Baptista, MZSP). COLOMBIA Cundinamarca: 
Sabana de Bogota, 4°43'N, 74°10'W, 10.xii.l990 (C. 
Valderrama, CV). ECUADOR Guaijas: Guayaquil, 
2°10'S, 79°.54'W, 18.iii.l942 (H. E. F. & D. E. F, 
CAS), 22.iii.1942 (Landis, CAS). Pichlncha: 1.5 mi N 
Quito, 0°0'N, 78°30'W, 23.ii.1955 (E. I. Sehlinger & 
E. S. Ross, CAS). Tungurahua: Baiios, 1°24'S, 
78°25'W, 15.ivI939 (W. C. Macintyre, MCZ); Puiiapi, 
1°22'S, 78°28'W, 19.vi.l943 (D. L. F. & H. E. F, 
CAS). PARAGUAY Alto Parana: Taguararaya [?], 
25°30'S, .54°50'W (AMNH). Caazapa: Villa Pastoreo, 
25°53'S, 55°45'W (D. Wees, MCZ). PERU Cajamar- 
ca: Cajamarca, 7°10'S, 78°31'W, 15.ii.l942 (W. Wey- 
rauch, AMNH). Lanihaijeque: 10 km S Chiclayo, 
7°59'S, 77°17'W, 19.iii.l951 (E. S. Ross & Michel- 
bacher, CAS). Piura: Cerro Prieto, La Brea, 4°41'S, 
81°6'W (CAS); Higueron (Las Lomas) [?], 4°19'S, 
80°26'W, 29.vii.194I (D. L. F & H. E. F, CAS). 
VENEZUELA Sucre: 1 km S Villa Frontado, Rd. to 



30 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Caripe, 10°27'N, 63°37'W, 12.ii.l984 (J. Coddington, 

USNM). 

7. Metepeira vigilax (Keyserling) 
Figures 36, 53-59, 327; Map 4 

Epeira vigilax Keyserling, 1893: 211, fig. 156, 6. 

Male holotype from Taquara, Rio Grande do Sul, 

Brazil, in BMNH, examined. 
Araneus vigilax: — Petrunkeviteh, 1911: 324. Roewer, 

1942: 856. 
Metepeira dominicana Archer, 1965: 132, figs. 12, 18, 

$ . Female holotype from west of Bani, Dominican 

Republic, in AMNH. Holotype lost. Brignoli, 1983: 

275. NEW SYNONYMY. 
Metepeira vigilax: — Levi, 1991: 180. Platnick, 1993: 

449. 

Note. Although the type for M. dominicana is 

lost, the name has been identified by using Archer's 

(1965) description and illustration. 

Description. Female from Trujillo, west 
of Bani, Dominican Republic. Carapace 
light around eyes with lateral posterior ex- 
tensions (Fig. 58). Legs dark, light rings on 
proximal ends of leg articles. Femur I with 
ro\v of four macrosetae on anterior side; 
three on anteroventral side. Dorsal folium 
darker than in most species; fleur-de-lis 
usually reduced to two white spots (Fig. 
58). Venter brownish gray with lighter 
margins. Wide, short, median white line 
with pair of white spots on either side of 
spiracle (Fig. 59). Sternum brownish black 
with wide, white line widening anteriorly 
(Fig. 59). Ratio of eye diameters: posterior 
inedians and anterior medians 1.2, anterior 
laterals 1.2, posterior laterals 1.1. Anterior 
median eyes separated by 1.8 diameters, 
posterior inedian eyes by 1.1, anterior me- 
dian eyes separated from anterior laterals 
by 3.5 diameters of anterior lateral eyes, 
lateral eyes separated by 0.1 their diame- 
ters. Total length 9.2 mm. Carapace 3.9 
mm long, 3.2 wide. First femur 4.1 mm, 
patella and tibia 4, metatarsus 3.3, tarsus 

1.2. Second patella and tibia 3.4 mm, third 

2.3, fourth 3.4. 

Male froin same locality as female. 
Black carapace with white around eyes and 
extending posteriorly; white wedge mark 
in center (Fig. 56). Legs ringed like fe- 
male. Femur I with row of three luacro- 
setae on anterior side; three on anterov- 



entral side. Dorsum and venter as in fe- 
male (Figs. 56, 57). Median white line may 
be limited to posterior end of sternum 
(Fig. 57). Ratio of eye diameters: posterior 
medians and anterior inedians 0.9, anterior 
laterals 1.1, posterior laterals 1. Anterior 
median eyes separated by 1.6 diameters, 
posterior median eyes by 1.3, anterior me- 
dian eyes separated from anterior laterals 
by 2 diameters of anterior lateral eyes, lat- 
eral eyes separated by 0.2 their diameters. 
Total length 3.9 mm. Carapace 2 mm long, 
1.5 wide. First femur 2.6 mm, patella and 
tibia 2.5, metatarsus 2.4, tarsus 0.9. Sec- 
ond patella and tibia 2.1 mm, third 1.2, 
fourth 1.7. 

Diagnosis. Female M. vigilax differ 
from those of other species in the M. vi- 
gilax species group by the shape of the 
epigynal openings: from a ventral view the 
openings are oval and angled inward pos- 
teriorly (Fig. 55); from a posterior view, 
the edges of the openings are more par- 
allel to the body (Fig. 61) as opposed to 
more pei^pendicular to the body (Figs. 39, 
61). Male M. vigilax differ from other spe- 
cies because the embolus is larger and 
more robust (compare Fig. 53 with Figs. 
38, 46, 60); the two flagella on the median 
apophysis are of more similar width (com- 
pare Fig. 53 with Fig. 38), and the keel on 
the median apophysis is slim and feather- 
shaped (Fig. 38), in contrast to arrow- 
shaped (Figs. 46, 60) or absent (Fig. 38). 
The dorsal folium differs from other Me- 
tepeira species by having a wide black me- 
dian stripe at the posterior end of the ab- 
domen (Fig. 58). In Brazilian and Bolivian 
specimens this stripe often extends all the 
way to the black anterior shoulders of the 
dorsum, forming a wide T-shape mark. 

Variation. Specimens from Argentina 
tend to be more lightly pigmented than 
those from more northern localities. White 
markings on the eye region of Brazilian 
and Bolivian specimens surround only the 
lateral eyes, in contrast to those on His- 
paniolan specimens, which cover the en- 
tire eye region. 

Natural History. Mature adults have 



Metepeira • Piel 31 







cajabamba 

(5) 




Figures 38^5. Metepeira cajabamba new species (sp. 5 [38-44] 7°37'S, 78°3'W; [45] 9°10'S, 77°45'W). 38, male palpus, 

mesal. 39, epigynum, posterior. 40, epigynum, ventral. 41 , male, dorsal. 42, male, ventral. 43, female, dorsal. 44, female, ventral. 

45, female, ventral. 

Figures 46-52. Metepeira glomerabilis (Keyserling) (sp. 6; 1°22'S, 78°28'W). 46, male palpus, mesal. 47, epigynum, posterior. 

48, epigynum, ventral. 49, male, dorsal. 50, male, ventral. 51, female, dorsal. 52, female, ventral. 

Figures 53-59. Metepeira vigilax (Keyserling) (sp. 7 [53-57] 18°27'N, 72°17'W; [58,59] 18°17'N, 70°22'W). 53, male palpus, 

mesal. 54, epigynum, posterior. 55, epigynum, ventral. 56, male, dorsal. 57, male, ventral. 58, female, dorsal. 59, female, ventral. 

Scale bars: dorsum and venter figures 1 .0 mm. 



32 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



been collected throughout the year except 
Januaiy, February, and March (Fig. 327). 
Spiders have been found among electric 
wires four meters above ground. Locality 
elevations range from near sea level to 
1,400 m and follow an ecological zone that 
decreases in elevation with distance from 
the equator (Fig. 36). Over equivalent lat- 
itudes, M. vigilax lives at less than one- 
tenth of the elevation of M. rectangula 
(Fig. 36). 

Distribution. Hispaniola, Bolivia, Brazil, 
and coastal Argentina (Map 4). The dis- 
junct distribution between Hispaniolan 
and South American populations may be 
due to human-assisted migration. 

Records Examined. ARGENTINA Buenos Aires: 
Celsya-Pereyra [?], 34°50'S, 58°6'W (MACN); Ze- 
laya, 34°21'S, 58°52'W (MACN). BOLIVIA La Paz: 
Apolo, 14°43'S, 68°31'W, 10.viii.l989 (L. E. Pena, 
AMNH). Santa Cruz: Estacion Robore, above creek, 
18°20'S, 59°45'W, 27.ix.1955 (Azambuya, CAS). 
BRAZIL Espirito Santo: Fazenda Santa Maria [?], 
Apiaca, 21°4'S, 41°25'W, 14.V.1988 (R. L. C. Baptista, 
MZSP). Rio Grande do Sul: Sao Leopoldo, 29°46'S, 
5r9'W, 14.vi.l964 (Celia Valle, MZSP). DOMINI- 
CAN REPUBLIC Aziia: El Pueriio, Majagual and 
Peralta, I8°34'N, 70°47'W, 10.xi.l979 (E. Marcano, 
MNSD). Prov. Tnijillo Vdldez: W Bani, 18°17'N, 
70°22'W, 8.viii.l958 (A. F. Archer & E. de Boyrie 
Moya, AMNH). HAITI Departement de L' Quest: 
Kenscoff, 18°27'N, 72°17'W, 15.xii.l929 (J. C. Myers, 
AMNH), 17.iv.l935 (AMNH). 

8. Metepeira rectangula (Nicolet) 
Figures 36, 60-66, 306; Map 4 

Epeira rectangula Nicolet, 1849: 500, female holo- 

type from Valdivia, Chile, in MNHN. 
Metepeira labyrinthea: — Petrunkevitch, 1911: 298. 

Roewer, 1942: 868. Bonnet, 1957: 2821. Erroneous 

synonymy. 
Metepeira rectangulata: — Chamberlin and Ivie, 1942: 

71. Unjustified emendation. 

Note. The name was identified using drawings of 

the holotype (H. W. Levi, personal illustrations). 

Description. Female from Angol, Mal- 
leco, Chile. Carapace reddish brown with 
long white setae behind lateral eyes. An- 
terior third of carapace white, median 
white line reaching thoracic furrow (Fig. 
65). Proximal halves of femora white, re- 
mainder black with distal white marks on 
dorsal surfaces. Patellae inostly black, re- 



maining articles white with black marks at 
base of setae. Femur I with row of three 
to five macrosetae on anterior side; three 
to four on anteroventral side. Anterior 
margin of dorsal abdomen black; dorsal fo- 
lium yellowish with brown speckles, white 
fleur-de-lis pattern with wide branches 
(Fig. 65). Venter black with wide white 
median line, flanked by pair of thin white 
lines; pair of white spots on either side of 
spiracle connected by white line (Fig. 66). 
Sternum black with posterior white mark 
(Fig. 66). Batio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.1, posterior laterals 1.1. Anterior 
median eyes separated by 1.7 diameters, 
posterior median eyes by 1.0, anterior me- 
dian eyes separated from anterior laterals 
by 3.3 diameters of anterior lateral eyes, 
lateral eyes separated by 0.1 their diame- 
ters. Total length 8.4 mm. Carapace 3.7 
mm long, 2.9 wide. First femur 4.2 mm, 
patella and tibia 4.1, metatarsus 3.7, tarsus 
1.2. Second patella and tibia 3.4 mm, third 

2.1, fourth 3.1. 

Male from Angol, Malleco, Chile. Car- 
apace reddish brown, anterior third white, 
median white line extending to thoracic 
furrow (Fig. 63). Proximal halves of fem- 
ora, white, distal halves black. Patellae 
black, remaining articles white widi black 
spots at base of setae. Feinur I with row 
of six to eight macrosetae on anterior side; 
seven to 11 on anteroventral side. Dorsal 
abdomen white, marbled, and speckled 
brown (Fig. 63). Venter dark brown with 
wide white median mark, flanked by pair 
of thin white lines; pair of white spots on 
either side of spiracle connected by white 
line (Fig. 64). Sternum dark brown with 
partly broken median white line (Fig. 64). 
Batio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 

1.2, posterior laterals 1.1. Anterior median 
eyes separated by 1.7 diameters, posterior 
median eyes by 1.1, anterior median eyes 
separated from anterior laterals by 2.9 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 6.1 mm. Carapace 3.2 mm long. 



Metepeira • Piel 



2.5 wide. First femur 5.2 mm, patella and 
tibia 5, metatarsus 4.9, tarsus 1.2. Second 
patella and tibia 4.1 mm, third 2, fourth 
3.1. 

Diagnosis. Female M. rectangula differ 
from the other species in the M. vigilax 
group (M. vigilax, M. glomerahilis, M. ca- 
jahamba) by the shape of the epigynal 
openings: from a ventral view they are 
larger, wider, and more gaping than in the 
other species (compare Fig. 62 with Figs. 
40, 48, 55). Males differ from other spe- 
cies in the group by having an embolus 
that is relatively larger than that of M. 
glomerahilis and M. cajabaniba, yet small- 
er than that of M. vigilax (compare Fig. 60 
with Figs. 38, 46, 53). Among all members 
of the M. vigilax group, the keel on the 
median apophysis of M. rectangula is the 
largest and most robust, and the dorsal fo- 
lium has the lightest coloration (Fig. 65). 

Va nation. Average body length of 11 fe- 
males examined 8.6 mm, range 5.8 to 10 
mm. Average body length of six males ex- 
amined 6 mm, range 4.1 to 7 mm. Speci- 
mens from two localities in western Ar- 
gentina resemble M. vigilax, and may be 
hybrids. 

Natural History. This species appears to 
follow a narrow ecological zone that de- 
creases in elevation with increasing south- 
ern latitude (Fig. 36). Median elevation, 
about 500 m. Mature specimens have 
been collected January through April (Fig. 
306). Specimens from localities south of 
the 36th parallel tend to be found in Jan- 
uaiy and Februaiy, whereas those north of 
the 36th parallel tend to be found in 
March and April. 

Distribution. Chilean Andes between 
31° and 38° south (Map 4). 

Records Examined. ARGENTINA Cordoba: Cala- 
muchita, 32°4'S, 64°33'W, 15.iii.l954 (J. M. Viana, 
MACN). Mendoza: Mendoza, 32°53'S, 68°49'W, 
30.iii.l965 (H. W. Levi, MCZ). CHILE Bio-Bio: 4 km 
E road to Pinto, 36°42'S, 71°53'W, 4.i.l976 (B. Mo- 
reno, AMNH); Road to Pemuco, Ci-uce del Carmen, 
36°56'S, 72°4'W, 10.i.l976 (G. Moreno, AMNH). Co- 
qnimbo: lUapel: Salamanca: Fundo Tahuinco, 
31°44'S, 71°5'W, 30.iv.l946 (R. Doneso, AMNH). 
Malleco: Angol, 37°48'S, 72°43'W (D. S. Bullock, 



CAS), 10.iii.l945 (E. A. Chapin, USNM). Maule: 10 
km S Curico, 35°4'S, 71°14'W, 15.iii.l968 (L. E. 
Peiia, MCZ); Cordillera de Parral [?], 36°9'S, 
71°50'W, 25.ii.1956 (L. E. Pena, IRSNB); Linares, 
35°51'S, 71°36'W (L. E. Peiia, IRSNB); Miraflores, 
Pedag. [?]. 35°55'S, 71°39'W (Toro, AMNH). 
O'Higgins: Fundo Millahue, Cunaco, 34°36'S, 
71°16'W, 30.iv.l961 (AMNH). Region Metropolitana: 
Melipilla, 33°42'S, 71°13'W (L. E. Pena, IRSNB). 
Valparaiso: Casablanca, 33°19'S, 71°25'W, 15.ii.l955 
(Edwdn Reed, AMNH). 



Metepeira labyrinthea Group 

Levi (1977) described the M. labyrin- 
thea group veiy broadly — it included spe- 
cies with a longitudinal white line down 
the sternum and a short keel on the me- 
dian apophysis. Here, this species group is 
much narrowed to include only three 
North American species: Metepeira laby- 
rinthea, Metepeira lacandon, and Metepei- 
ra spinipes. Males of these three species 
are unique among Metepeira by having a 
toothless, smooth keel on the median 
apophysis. In addition, their distal embolic 
apophysis rises away (anteriorly) from the 
embolus proper and projects forward (ven- 
trally) until it is almost even with the em- 
bolus tip (Figs. 67, 76). In contrast, other 
Metepeira species with distal embolic 
apophyses have the embolus tip extend far 
beyond the projection of the apophysis 
(e.g., Fig. 171). The female epigynum has 
a characteristic shape. The scape is thick 
and fleshy and the epigynal openings have 
membranes that make them look distinctly 
(Fig. 69) or indistinctly (Fig. 78) almond- 
shaped. The epigyna of the M. labyrinthea 
species group are easily confused with that 
of the closely related species, M. gosoga 
(Fig. 173). Although most differences be- 
tween M. labyinthea group and M. go,soga 
are only obvious in the epigynum, it 
should be noted that the dark marks inside 
the epigynal openings of the former ap- 
pear to look cross-eyed, but this cannot be 
said for the latter. In this work, only the 
species collected at localities south of the 
U.S. /Mexico boarder are treated. 



34 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



9. Metepeira spinipes 
F. O. P. -Cambridge 
Figures 67-75, 335; Map 6 

Metepeira spinipes F. O. P.-Cambridge, 1903: 459, 
figs. 9, 10, (J, ?. Male holotype from Mexico City, 
Mexico, in BMNH, examined. Roewer, 1942: 868. 

Epeira labyrinthea grinnelli Coolidge, 1910: 281, 9. 
Holotype from Palo Alto, California, lost. 

Araneus spinipes: — Petrunkevitch, 1911: 317. 

Aranea labyrinthea grinnelli: — Moles, 1921: 42. 

Metepeira douglasi Chamberlin and Ivie, 1941: 18, 
figs. 21-23, 9 . Female holotype from Santa Ana, 
California, in AMNH, examined. Chamberlin and 
Ivie, 1942: 66, figs. 169-170. First synonymized 
with M. labyrinthea grinnelli by Levi, 1977: 198. 

Metepeira labyrinthea grinnelli: — Roewer, 1942: 868. 

Metepeira labyrinthea: — Bonnet, 1957: 2822. Erro- 
neous synonyiTiy. 

Metepeira grinnelli: — Levi, 1977: 198, figs. 21-27, S , 
? . NEW SYNONYMY. 

Description. Female from Huitzilac, 
Morelos, Mexico. Brown carapace with an- 
terior portion darker reddish brown, white 
behind lateral eyes (Fig. 74). Legs yellow- 
ish, femora reddish brown distally, otlier 
articles dark brown distally. Femur I with 
row of four to five macrosetae on anterior 
side; two to seven on anteroventral side. 
Anterior shoulders of abdomen black. 
Dorsal folium with usual Metepeira pat- 
tern, though largest branches of white 
fleur-de-lis shape usually widened into 
large spots (Fig. 74). Venter of abdomen 
black with wide white median line (Fig. 
75). Pair of small white spots on either side 
of spiracle. Sternum black with posterior 
white mark that in some cases extends an- 
teriorly to the labium (Figs. 73, 75). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.1, anterior laterals 1.0, 
posterior laterals 0.9. Anterior median 
eyes separated by 1.9 diameters, posterior 
median eyes by 1.3, anterior median eyes 
separated from anterior laterals by 2.9 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 9.2 mm. Carapace 3.7 mm long, 
2.9 wide. First femur 4.4 mm, patella and 
tibia 4.7, metatarsus 4.6, tarsus 1.3. Sec- 
ond patella and tibia 4.1 mm, third 2.4, 
fourth 3.6. 

Male from Huitzilac, Morelos, Mexico. 



Carapace, dorsum, venter, sternum as in 
female (Figs. 71, 72). Distal portions of leg 
articles reddish black, elsewhere yellowish. 
Femur I with row of four to six macrosetae 
on anterior side; five to nine on anterov- 
entral side. Ratio of eye diameters: poste- 
rior medians and anterior medians 1.1, an- 
terior laterals 1.2, posterior laterals 1.1. 
Anterior median eyes separated by 1.8 di- 
ameters, posterior median eyes by 1.1, an- 
terior median eyes separated from anterior 
laterals by 3.2 diameters of anterior lateral 
eyes, lateral eyes separated by 0.3 their di- 
ameters. Total length 7.5 mm. Carapace 
3.8 mm long, 2.8 wide. First femur 6.2 
mm, patella and tibia 6.3, metatarsus 7.1, 
tarsus 1.8. Second patella and tibia 5.3 
mm, third 2.5, fourth 4. 

Diagnosis. Inside each epigynal depres- 
sion of M. spinipes and M. lacandon is a 
membrane that forms a slanted, oval- 
shaped opening (Figs. 69, 78). Within each 
oval-shaped opening is a dark mark, which 
in M. spinipes takes up a small part of that 
opening, resulting in a cross-eyed appear- 
ance (Fig. 69). Also, in M. spinipes (Fig. 
69), the edges of the epigynal depressions 
are less distinct than in M. lacandon (Fig. 
78). The distal embolic apophysis of M. 
spinipes is arrow-shaped (Fig. 79), in con- 
trast to a wider shovel-shape in M. lacanA 
don (Fig. 79). 

Variation. Average body length of 54 fe- 
males examined 9.1 mm, range 5.5 to 12.4 
mm. Average body length of 55 males ex- 
amined 6.6 mm, range 3 to 10.8 mm. 

Natural History. Mature specimens 
have been collected between August and 
early November (Fig. 335). Elevations 
range from near sea level in California to 
2,600 m in central Mexico. Variation in 
sexual dimoi-phism appears to correlate 
with habitat and social structure (Piel, 
1996). Webs are found in dry regions 
among mesquite, Opuntia, Agave (ma- 
guey), cultivated Yucca, and Cactus. Spi- 
ders live in medium to small social colo- 
nies, which vary in size in accordance with 
local habitat quality (Uetz, 1988a,b). This 
behavioral and ecological relationship is 



Metepeira • Piel 35 



4 M 




82 

lacandon 
(10) 



Figures 60-66. Metepeira rectangula (Nicolet) (sp. 8; 37°48'S, 72°43'W). 60, male palpus, mesal. 61, epigynum, posterior. 62, 

epigynum, ventral. 63, male, dorsal. 64, male, ventral. 65, female, dorsal. 66, female, ventral. 

Figures 67-75. Metepeira spinipes F. O. P. -Cambridge (sp. 9 [67-72,74,75] 19'=0'29"N, 99°15'50"W; [73] 39°18'N, 123°48'W). 

67, male palpus, mesal. 68, epigynum, posterior. 69, epigynum, ventral. 70, male embolic division, ventral. 71, male, dorsal. 72, 

male, ventral. 73, female, ventral. 74, female, dorsal. 75, female, ventral. 

Figures 76-83. Metepeira lacandon new species (sp. 10; 16°45'N, 92°38'W). 76, male palpus, mesal. 77, epigynum, posterior. 

78, epigynum, ventral. 79, male embolic division, ventral. 80, male, dorsal. 81, male, ventral. 82, female, dorsal. 83, female, 

ventral. 

Scale bars: dorsum and venter figures 1.0 mm. 



36 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



thought to occur as a result of the spiders 
pursuing a risk-sensitive foraging strategy 
(Uetz, 1996). 

Distribution. Oregon to central Mexico 
(Levi, 1977, map 1; Map 6). 

Records Examined. MEXICO Agiiascalientes: Hwy 
45, 5.3 mi N Aguascalientes, 21°57'N, 102°17'W, 
7.ix.l967 (R. E. Leech, REL). Baja California Sun 
Sierra Laguna, 17 air mi ENE Todos Santos, 
23°34'N, 110°0'W, 15.xii.l979 (C. E. Griswold, CAS). 
Chihuahua: 22.4 mi S Miiaaca, 28°24'N, 107°26'W, 
23.viii.1950 (R. Smith, AMNH). Durango: 10 mi E 
El Salto, 23°12'N, 105°52'W, 8.viii.l947 (W. J. 
Gertsch, AMNH); 11 km W Suehil, 23°35'N, 
104°5'W, 5.ix.l984 (W. J. Pulawsld, CAS); 6 mi NE 
El Salto, 23°19'N, 105°50'W, ll.viii.l947 (W. J. 
Gertsch, AMNH); Las Puentes [?], 26°49'N, 
106°2'W, 23.vii.1947 (W. J. Gertsch, AMNH); Oti- 
napa, 24°11'N, 105°2'W, 12.viii.l947 (W. J. Gertsch, 
AMNH); Palos Colorados, 24°2'N, 104°54'W, 
5.viii.l947 (W. J. Gertsch, AMNH); Providencia. 
26°44'N, 105°56'W, 24.viii.1947 (A. M. Davis, 
AMNH); SW Durango, 23°59'17"N, 104°45'47'W, 
22.X.1994 (W. H. Piel, MCZ). Guanajuato: 30 mi SE 
Leon, 6 mi SE Silao, 20°52'N, 101°21'W, 6.ix.l964 
(Jean & Wilton Ivie, AMNH); 6.3 mi NW Leon, 
21°13'N, 101°43'W, 6.ix.l967 (R. E. Leech, REL); 
between Moroleon & Cuitzeo, 20°5'36"N, 
101°9'28'W, 20.X.1994 (W. H. Piel, MCZ); near San 
Miguel de Allende, 20°55'N, 100°45'W, 16.lx.1976 
(C. E. Grisv^^old, CAS); S San Miguel de Allende, 
20°46'36"N, 100°47'28"W, 20.x. 1994 (W. H. Piel, 
MCZ); San Miguel de Allende. Road SW town, 
20°52'N, 100°56'W 25.x. 1982 (George Uetz, MCZ). 
Hidalgo: 18 mi E Huichapan, off Hwy 45, 20°23'N, 
99°22'W, 25.viii.1984 (W D. Sissom, C. Myers & L. 
Bom, MCZ); 4 mi N Tizayuca, 19°54'N, 98°59'W 
20..xi.l946 (E. S. Ross, CAS); 41 km N Zimapan, 
20°54'N, 99°13'W, 10.viii.l991 (W H. Piel & G. S. 
Bodner, MCZ); Apulco, 20°19'N, 98°20'W, 6.X.1947 
(H. Wagner, AMNH); Ozumbilla, 20°9'N, 101°16'W, 
2.X.1957 (R. Dreisbach, MCZ); Pachuca, 20°7'N, 
98°44'W, 30.viii.l957 (R. Dreisbach, MCZ); Tenango 
de Doria, 20°19'N, 98°13'W, 5.x. 1947 (H. Wagner, 
AMNH). Jalisco: 12 mi S Mazamitla, 19°47'N, 
103°8'W, 5.xii.l948 (H. B. Leech, CAS); Charco 
Ondo, 30 km W Ojuelos, 21°47'N, 101°53'W 
25. ix. 1945 (H. Wagner, AMNH). Mexico: Ixtapan de 
la Sal, 18°50'N, 99°41'W, 24.viu.1946 (H. Wagner, 
AMNH); Nevado de Toluca, 19°18'N, 99°44'W, 
8.iv.l979 (George Uetz, MCZ); San Juan Teotihu- 
acan, 19°41'N, 98°52'W, 4..xi.l939 (C. M. Bogert & 
H. E. Vokes, AMNH); Tenancingo, 18°58'N, 
99°36'W, 6.LX.1946 (H. Wagner, AMNH), l.x.1946 
(H. Wagner, AMNH), 15.x. 1946 (H. Wagner, 
AMNH); Tenango del Valle, 19°7'N, '99°33'W, 
25.viii.1946 (H. Wagner, AMNH), 27.viii.1946 (H. 
Wagner, AMNH); Teotihuacan, 19°41'N, 98°52'W 
31.viii.l959 (A. F. Ai-cher, AMNH); Tepotzotlan, 



19°43'N, 99°13'W, 26.X.1982 (George Uetz, MCZ), 
21. ii. 1983 (George Uetz, MCZ), 5.x. 1983 (George 
Uetz, MCZ); Toluca, 19°18'N, 99°44'W, 10.viii.l978 
(George Uetz, MCZ), l.viii.l986 (George Uetz, 
MCZ); Toluca, at bottom of mountain near Parque 
Cierra Morelos, 19°18'N, 99°44'W, 10.viii.l978 
(George Uetz, MCZ); Toluca, E of town on Paseo 
ToUocan [?], 19°18'N, 99°42'W, 23.X.1982 (George 
Uetz, MCZ). Mexico D. F.: 19°25'N, 99°10'W, 
12.X.1940 (H. Wagner, AMNH), 28.xii.1940 (R. H. 
Crandall, AMNH), 15.ix.l943 (H. & D. Frizzell, 
AMNH); Contreras, 19°18'N, 99°17'W, 4.xii.l944 (H. 
Wagner, AMNH), 15.ix.l965 (N. L. H. Krauss, 
AMNH); Delegacion Tlalpan, Colonia Santa Ursula 
Xitla, 19°16'0"N, 99°10'25'W, 12.x. 1994 (W H. Piel, 
MCZ); Desierto de los Leones, 19°22'N, 99°16'W, 
15.ix.l941 (H. Wagner, AMNH); El Xitle, 18°61'N, 
99°17'W [?], 12.viii.l942 (C. Tellez, AMNH); Haci- 
enda Cordoba, 19°26'N, 99°10'W [?], 29.X.1944 (H. 
Wagner, AMNH); Ouieros, 18°62'N, 99°17'W [?], 
5.vii.l943 (M. Cardenas, AMNH); Mexico City, 
19°25'N, 99°10'W, l.xi.l941 (C. Velo, AMNH), 
25.Lx.1957 (R. Dreisbach, MCZ); Mixcoac, 19°23'N, 
99°11'W (AMNH), 13.X.1940 (A. F. Archer, AMNH); 
Mixenac, 19°25'N, 99°10'W 13.X.1940 (H. Wagner, 
AMNH); Pedregales, 18°60'N, 99°17'W [?], 
15.viii.l909 (AMNH); Petregal [?], 18°60'N, 
99°17'W l.xii.l943 (AMNH); Rancho Cordoba, 
19°27'N, 99°10'W, 29.X.1944 (H. Wagner, AMNH); 
Tlaplan, 19°17'N, 99°10'W, 7.viii.l991 (W. H. Piel & 
G. S. Bodner, MCZ). Michoacan: 25 mi W La Barca 
nr Lago de Chapala, 20°17'N, 102°34'W, ll.ix.l976 
(C. E. Griswold & Jackson, CAS); between Patzcuaro 
& Uruapan, 19°29'19"N, 101°48'20"W, 19.X.1994 (W 
H. Piel, MCZ); Hills N of Patzcuaro, 19°45'N, 
101°36'W, 24.viii.1959 (A. F. Archer, AMNH); Hwy 
110, 4 mi W. Jiquilpan, 19°59'N, 102°47'W, 
2.viii.l967 (R. E. Leech, REL); Hwy 15, 9.5 mi W 
Morelia, 19°42'N, 101°16'W; 18.viii.l967 (R. E. 
Leech, REL); Lake Chapala, NW of Cojumatlan, 
20°10'N, 102°53'W, 7.ix.l966 (Jean & Wilton Ivie, 
AMNH); Monte de Zacapu, 19°47'N, 101°50'W, 
24.viii.1959 (A. F. Archer, AMNH). Morelos: Cuer- 
navaca, 18°55'N, 99°15'W (AMNH), 15.ix.l941 (H. 
Wagner, AMNH), 18.xi.l946 (M. G. Bradt, AMNH); 
Cueniavaca/Tepotzotlan, interchange between 1-95 & 
115, 18°55'N, 99°13'W, 7.viii.l978 (MCZ); Huitzilac, 
19°2'2"N, 99°16'13'W^, 13.X.1994 (W. H. Piel, MCZ); 
North of Cuernavaca, 18°58'11"N, 99°14'37"W, 
11.x. 1994 (W. H. Piel, MCZ); S of Huitzilac, 
19°0'29"N, 99°15'50"\'V, 16.x. 1994 (W H. Piel, MCZ). 
Puehla: 6 mi E Rio Frio, 19°20'N, 98°35'W, 
22.viii.1964 (Jean & Wilton Ivie, AMNH); Puebla, 
19°3'N, 98°12'W, 21.x. 1982 (George Uetz, MCZ). 
San Luis Potosi: 3 km W Pilares, 21°55'34"N, 
100°48'6'W, 21.X.1994 (W. H. Piel, MCZ); Cuidaddel 
Maiz, 22°24'N, 99°36'W, 25.viii.1954 (R. Dreisbach, 
MCZ). Sonora: 46 mi S Agua Prieta on Highway 10, 
31°0'N, 109°16'W, 15.viii.l959 (B. A. Branson, 
AMNH); Hermosillo, 29°4'N, 110°55'W, 20.lx.1952 
(B. Malldn & V. E. Thatcher, AMNH); Sierra Man- 



Metepeira • Piel 



37 



zanal, 30°50'N, 110°10'W, 14.ix.l976 (Roth & 
Schroepfer, MCZ). Tlaxcala: Huamantla, 19°19'N, 
97°56'W [?], 15.vii.l981 (C. Gold, CAS). Veracruz: 
15 mi W. Banderilla, 19°39'N, 97°8'W, 31.X.1973 (S. 
C. Williams & C. L. Mullinex, CAS); 15 mi West of 
Jalapa, 19°32'N, 97°9'W, 23.vi.1946 (A. M. & L. I. 
Davis, AMNH). Zacatecas: 13 mi N. Sombrerete, 
23°44'32"N, 103°47'10"W 22.X.1994 (W H. Piel, 
MCZ); Canutillo, 24°47'N, 101°31'W, 14.viii.l947 (W. 
J. Gertsch, AMNH); S. Zacatecas, 22°45'7"N, 
102°29'37"W, 22.X.1994 (W. H. Piel, MCZ). USA. Ar- 
izona: Southwestern Research Station, Chiricaliua 
Mtns., 31°35'N, 109°14'W, 20.viii.l976 (V. Roth, 
MCZ). California: 26 mi W. Santa Rosa on Hwy 116, 
38°31'N, 123°4'W, 19.ix.l976 (M. E. Thompson, 
MCZ); Mendocino, 39°18'N, 123°48'W, 18.viii.l959 
(W J. Gertsch, MCZ); Monterey, 36°36'N, 121°54'W, 
l.ix.l949 (A. F. Archer, MCZ); Pacific Grove, 
36°37'N, 121°56'W (R. V. Chamberlin, MCZ); Palo 
Alto, 37°27'N, 122°9'W (Doane, MCZ); Salt marsh 
on N shore of San Pablo Bay, Vallejo, 38°8'N, 
122°27'W (D. Spiller, MCZ). 

10. Metepeira lacandon new species 
Figures 76-83, 332; Map 6 

Holotijpe. Male from San Cristobal, Chiapas, Mexico. 
The specific name is a noun in apposition after the 
Indian people who live in Chiapas. Holotype de- 
posited in the AMNH. 

Description. Female paratype from San 
Cristobal, Chiapas, Mexico. Reddish cara- 
pace, slightly darker in anterior half, ligh- 
ter behind lateral eyes (Fig. 82). Leg ar- 
ticles yellowish, gradually turning reddish 
brown distally. Femur I with row of four 
or five macrosetae on anterior side; two to 
four setae on anteroventral side. Anterior 
shoulders of abdomen black. Branches of 
white fleur-de-lis shape in dorsal folium 
thinner than in most species (Fig. 82). 
Venter of abdomen black with wide white 
median line that extends about half the 
distance between epigynal groove and 
spinnerets (Fig. 83). Pair of very small 
white spots on either side of spiracle. Ster- 
num black, often with central w^hite spot 
(Fig. 83). Ratio of eye diameters: posterior 
medians and anterior medians 1.0, anterior 
laterals 1.3, posterior laterals 1.2. Anterior 
median eyes separated by 1.7 diameters, 
posterior median eyes by 1.0, anterior me- 
dian eyes separated from anterior laterals 
by 4.5 diameters of anterior lateral eyes, 
lateral eyes separated by 0.1 their diame- 



ters. Total length 11.5 mm. Carapace 4.6 
mm long, 3.6 wide. First femur 5.3 mm, 
patella and tibia 5.3, metatarsus 4.9, tarsus 

1.6. Second patella and tibia 4.4 mm, third 

2.7, fourth 4. 

Male holotype. Carapace, dorsum, ven- 
ter, sternum as in female (Figs. 80, 81). 
Distal halves of femora, tibia reddish 
brown, elsewhere yellowish. Patellae, 
metatarsi reddish. Femur I with row of 
four or five macrosetae on anterior side; 
six or seven on anteroventral side. Ratio of 
eye diameters: posterior medians and an- 
terior medians 1.1, anterior laterals 1.5, 
posterior laterals 1.3. Anterior median 
eyes separated by 1.7 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 3.3 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 6.8 mm. Carapace 3.5 mm long, 
2.7 wide. First feinur 5.3 mm, patella and 
tibia 3.9, metatarsus 5.6, tarsus 1.7. Sec- 
ond patella and tibia 4.4 mm, third 2.3, 
fourth 3.4. 

Diagnosis. Inside each epigynal depres- 
sion of M. lacandon and M. spinipes is a 
membrane that forms a slanted, oval- 
shaped opening (Figs. 69, 78). Within each 
oval-shaped opening is a dark mark, which 
in M. lacandon takes up only a large part 
of that opening, resulting in a less cross- 
eyed appearance (Fig. 78). Also, in M. la- 
candon (Fig. 78), the edges of the epigynal 
depressions are luore distinct than in M. 
spinipes (Fig. 69). The distal embolic 
apophysis of M. lacandon is more shovel- 
shaped (Fig. 79) than the thinner, arrow- 
shaped one in M. .spinipes (Fig. 70). 

Variation. Average body length of three 
females examined 8.5 mm, range 7.5 to 10 
mm. Average body length of four males 
examined 5.8 mm, range 4.2 to 6.8 mm. 

Natural History. Mature specirnens 
were collected between July and Septem- 
ber (Fig. 332) from oak— pine woodland. 
Elevations range from 1,700 to 2,300 m. 

Distribution. Mountainous regions of 
Chiapas, Mexico (Map 6). 



38 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Records Examined. MEXICO Chiapas: 12 km NW 
Comitan, 16°23'N, 92°15'W, 30.\dii.l976 (E. S. Ross, 
CAS); 4 mi SE San Cristobal, 16°42'N, 92°36'W, 
23.viii.1966 (Jean & Wilton Ivie, AMNH); 5 km W 
San Cristobal de Las Casas on HWY 190, 16°44'N, 
92°41'W, 27.vii.1983 (W. Maddison & R. S. Anderson, 
MCZ); 5 mi W San Cristobal, 16°45'N, 92°41'W, 
24.viii.1966 (Jean & Wilton Ivie, AMNH); San Cris- 
tobal, 16°45'N, 92°38'W, 13.ix.l947 (H. Wagner, 
AMNH); San Cristobal de las Casas, 16°45'N, 
92°38'W, 22.vii.1947 (C. J. & M. Goodnight, 
AMNH); Tenejapa, 16°49'N, 92°31'W, 22.vii.1950 (C. 
J. & M. Goodnight, AMNH). 

Metepeira nigriventris Group 

There are five species in the Metepeira 
nigriventris group: Metepeira nigriventris, 
Metepeira tarapaca, Metepeira calamuchi- 
ta, Metepeira galatheae, and Metepeira 
karkii. These closely related species are of- 
ten hard to distinguish because their gen- 
italia are similar, yet highly variable within 
a species. This species group is easily rec- 
ognized by the distinctive shape of the 
scape and similarities in palp moiphology. 
Typically the base of the scape originates 
anteriorly and projects ventrally before 
curving posteriorly. This projection creates 
an overhang and a noticeable gap between 
the scape and the genital openings (e.g., 
Figs. 85, 86, 101, 102). The embolus is 
thick and has a large, prominent, distal 
apophysis that hides under the terminal 
apophysis (Figs. 84, 92, 100, 110, 121). 

1 1 . Metepeira nigriventris (Taczanowski) 
Figures, 84-91, 310; Map 5 

Epeira nigriventris Taczanowski, 1878: 151, fig. 6, ?. 
Female lectotype from Lake Junin, Peru, in PAN, 
type lost. Keyserling, 1893: 217, fig. 161, 9,3. 

Araneus nigriventris: — Chamberlin, 1916: 248. Bon- 
net, 1955: 550. 

Metepeira nigriventris: — Chamberlin and Ivie, 1942: 
74, figs. 211-214, 9,3. Platnick, 1993: 449. 

Note. Although the type is lost, the type locality 
and Taczanowski s descriptions are sufficient to rec- 
ognize the species. 

Description. Female from 12 km west of 
Tarma, Junin, Peru. Carapace dark brown, 
light around eyes with lateral posterior ex- 
tensions (Fig. 90). Proximal halves of leg 
articles yellow, distal halves black. Femur 
I with row of four macrosetae on anterior 



side; five on anteroventral side. Dorsum 
darker and white fleur-de-lis pattern small- 
er than in most species (Fig. 90). Venter 
mostly black with reduced, short, thin, 
white median line (Fig. 91). SternuiTi en- 
tirely black. Ratio of eye diameters: pos- 
terior medians and anterior medians 1.0, 
anterior laterals 1.4, posterior laterals 1.4. 
Anterior median eyes separated by 1.8 di- 
ameters, posterior median eyes by 1.2, an- 
terior median eyes separated from anterior 
laterals by 5 diameters of anterior lateral 
eyes, lateral eyes separated by 0.4 their di- 
ameters. Total length 11 mm. Carapace 5.1 
mm long, 4.2 wide. First femur 5.5 mm, 
patella and tibia 5.8, metatarsus 5.5, tarsus 
1.9. Second patella and tibia 5.3 mm, third 
3.3, fourth 4.5. 

Male from same locality as female. An- 
terior margin of chelicerae with large, 
swollen tooth and several denticles. Cara- 
pace reddish brown with lighter eye re- 
gion, lateral posterior extensions, and long, 
thin median white line (Fig. 88). Femur 1 
with row of about five macrosetae on an- 
terior side; nine on anteroventral side. 
Coloration of legs, dorsum, venter, and 
sternum as in female (Figs. 88, 89). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.1, anterior laterals 1.3, 
posterior laterals 1.3. Anterior median 
eyes separated by 1.8 diameters, posterior 
median eyes by 1.2, anterior median eyes 
separated from anterior laterals by 3.7 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.4 their diameters. To- 
tal length 9 mm. Carapace 4 mm long, 3.2 
wide. First femur 5.8 mm, patella and tibia 
6.2, metatarsus 6.5, tarsus 1.7. Second pa- 
tella and tibia 4.5 mm, third 2.8, fourth 4. 

Diagnosis. Metepeira nigriventris is eas- 
ily distinguished from other species in the 
M. nigriventris group by its dark pigmen- 
tation. As its name implies, the sternum is 
black and the white ventral mark on the 
abdomen is reduced to a much shorter and 
thinner line (Figs. 88-91). While the ster- 
num of M. karkii is similarly dark, the dor- 
sal and ventral markings on the abdomen 
are much lighter (compare Figs. 88-91 



Metepeira • Piel 39 





nigriventris 
(11) 



i9& 9^^^ 99 

« tarapaca 



(12) 



A. iBi 




H 106 107 _ 

calamuchita 
105 - (13) 



Figures 84-91. Metepeira nigriventris (Taczanowski) (sp. 11; ir25'S, 75°48'W). 84, male palpus, mesal. 85, epigynum, pos- 
terior. 86, epigynum, ventral. 87, male embolic division, ventral. 88, male, dorsal. 89, male, ventral. 90, female, dorsal. 91, 

female, ventral. , „„ . 

Figures 92-99 Metepeira tarapaca new species (sp. 12; 21°39'S, 69°33'W). 92, male palpus, mesal. 93, epigynum, postenor. 

94, epigynum, ventral. 95, male embolic division, ventral. 96, male, dorsal. 97, male, ventral. 98, female, dorsal. 99, female, 

ventral. , ,.,„.,• 

Figures 100-107. Metepeira calamuchita new species (sp, 13; 32°4'S, 64°33'W). 100, male palpus, mesal. 101, epigynurri, 
posterior. 102, epigynum, ventral. 103, male embolic division, ventral. 104, male, dorsal. 105, male, ventral. 106, female, dorsal. 
107, female, ventral. 
Scale bars: dorsum and venter figures 1.0 mm. 



40 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



with Figs. 125-128). The epigynum of M. 
nigriventris has a wider membrane over 
the openings that is visible on either side 
of a very wide scape (Fig. 86). In contrast, 
this feature is hidden behind a thinner 
scape in its Hkely sister species, M. tara- 
paca (Fig. 94). The overall shape of tlie 
epigynum (Fig. 86) and the pair of small 
notches in the posterior lobes (Fig. 85) 
also make this species distinctive. The 
male palp of M. nigriventris has an em- 
bolus that is relatively sliinmer and more 
graceful than those of other species in the 
M. nigriventris species group (compare 
Fig. 84 with Figs. 92, 100, 109-110, 121). 
The shape of the embolus and its distal 
apophysis differs from other species (com- 
pare Fig. 87 with Figs. 95, 103, 112, 124). 

Variation. Average body length of 25 fe- 
males examined 9.5 mm, range 7.2 to 11.5 
mm. Average body length of 16 males ex- 
amined 7.4 mm, range 5.3 to 9.5 mm. 

Natural History. Spiders are commonly 
found around Lake Titicaca living in me- 
dium and large colonies among power 
lines, Bolivian pines. Cactus, Bacharis, 
rock outcroppings, and tall grasses (L. 
Rayor, personal communication, and vari- 
ous locality labels). Mature speciinens 
have been collected throughout the year 
except September and October (Fig. 310). 
Median elevation, 3,900 m. 

Distribution. High altitude regions of 
southern Peru and western Bolivia (Map 
5). 

Records Examined. ARGENTINA Jujuy: Puma- 
huasi, 22°17'S, 65°41'W, 8.xi.l970 (L. E. Pena, 
MCZ). BOLIVIA La Paz: 45 mi S La Paz, 17°9'S, 
67°36'W, 25.ii.1951 (E. S. Ross & Michelbacher, 
CAS); 70 mi S La Paz, 17°30'S, 67°36'W, 25.ii.1951 
(E. S. Ross & Michelbacher, CAS); La Paz, Avenida 
Sport Club, 16°30'S, 68°9'W, 4.i.l959 (A. M. Nadler, 
AMNH); La Paz, in garden of house, 16°30'S, 
68°9'W, 15.iv.l959 (R. Walsh, AMNH); Lake Titicaca, 
Copacabana, Yampupata, & Isla del Sol, 16°10'S, 
69°5'W, 17.V.1995 (L. Rayor, MCZ); near La Paz, 
16°30'S, 68°9'W, 24.V.1958 (R. Walsh, AMNH); S end 
of Lake Titicaca, 100 km NW La Paz, 16°10'S, 
69°5'W, 5.vii.l958 (R. Walsh, AMNH); Tialiuanaco, 
Puma Puerto Ruins, 16°33'S, 68°42'W, l.ii.l973 (Ann 
Moreton, MCZ). Oruro: 6 km N Challapata, 18°51'S, 
66°47'W, 23.ii.1951 (E. S. Ross & Michelbacher, 



CAS); Gorge Uhuschlucht, near Oruro, 17°59'S, 
67°9'W, 7.ii.l954 (Forster & Schindler, ZSM). Potosi: 
Villazon, 22°6'S, 65°36'W, 30.xii.l984 (L. E. Peiia, 
AMNH). PERU Apurimac: Chincheros, 13°30'48"S, 
73°42'47"W, 12.xii.l980 (C. Gold, CAS); Puna near 
Abancay, 13°38'2"S, 72°52'52"W, 15.xii.l947 (W. Wey- 
rauch, CAS). Ayacucho: Puquio, 14°42'S, 74°8'W, 
15. iv. 1950 (F. Blancas, MUSM); San Antonio (Pu- 
quio), 14°47'S, 74°7'W, l.xi.l985 (D. Silva, MUSM). 
Cusco: Cheqquerec, 13°23'S, 72°8'W, 2.ix.l993 (J. 
Ochoa Camara, MCZ); Cusco, 13°31'6"S, 
71°58'41"W, 8.viii.l965 (P & B. Wygodzinsky, 
AMNH). junin: 8 mi W Tarma, 11°25'S, 75°48'W, 
6.i.l955 (E. I. Schlinger & E. S. Ross, CAS); Cochas 
Bajo, 11 km W Tarma, 11°25'21"S, 75°46'11"W, 
27iii.l988 (J. Palmer & D. Smith, MCZ); Cochas 
Bajo, 11 km W Tarma, rock ledge in agricultural val- 
ley, 11°25'21"S, 75°46'11"W, 29.iii.1988 (J. Palmer, 
MCZ); Huancayo, 12°4'S, 75°14'W, 15.vi.l947 (W. 
Weyrauch, AMNH); Oroya, 11°32'S, 75°54'W, 
12.iv.l914 (M. P. Anderson, AMNH). Lima: Bosque 
de Zarate, 11°53'S, 76°27'W, 18.i.l981 (J. Francke, 
MUSM). Puno: 10 mi N Ayaviri, 14°45'S, 70°35'W, 
l.iii.l951 (E. S. Ross & Michelbacher, CAS); Cama- 
cane, 15°55'S, 69°50'W, 20.xi.l955 (L. E. Peiia, 
IRSNB); Isla Taquih, Lago Titicaca, 15°46'S, 
69°41'W, 23.xii.1980 (C. Gold, CAS); Juh (col. Chu- 
cuito), 16°13'S, 69°27'W, 7.xi.l952 (F. Blancas, 
MUSM); near Chucuito, Lago Titicaca, 15°50'S, 
69°48'W, 10.iii.l953 (M. Koepcke, MUSM); Puna, 
Lake Titicaca, 15°50'S, 70°2'W, 15.vi.l947 (W Wey- 
rauch, AMNH); Puno, 15°50'S, 70°2'W (Soukup, 
AMNH); Yunguyo, downtown plaza, 16°15'S, 69°5'W, 
31.i.l973 (Ann Moreton, MCZ). 

12. Metepeira tarapaca new species 
Figures 92-99, 305; Map 7 | 

Holotype. Male from Quillagua, Antofagasta, Chile, 
4.ii.l965, L. E. Pena, in MCZ. The specific name 
is a noun in apposition after a Chilean province 
where it is abundant. 

Description. Female paratype from 
Quillagua, Antofagasta, Chile. Light red- 
dish brown carapace, white in center and 
around eyes with lateral posterior exten- 
sions (Fig. 98). Legs yellowish white, 
ringed brown at distal ends of articles. Fe- 
mur I with row of four macrosetae on an- 
terior side; three on anteroventral side. 
Dorsal folium white with black speckles 
(Fig. 98). Venter of abdomen black with 
wide, white median line, sometimes 
flanked by thinner white lines that togeth- 
er form a U-shape posteriorly (Fig. 98). 
Sternum black with median white line, 
sometimes broken (Fig. 99). Ratio of eye 



Metepeira • Piel 41 



4500j 
400O- 
350O- 
3000-- 
250a- 

20oa- 

1500- - 

1000-- 

500- - 

.- 



cP o ° 







20 40 60 80 100 

Percent of median white line on sternum 



Figure 108. Elevation of collection localities of mature female 
M. tarapaca with differing amounts of whiite on the sternum. 
Spiders with a median white line covering 100% of the sternum 
length are found at a wide altitude range. Spiders with a me- 
dian white mark covering only a short length of the sternum 
are only found at high elevations. Elevations estimated from 
NOAA database of 5- by 5-minute geographic tiles. 



diameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.3, posterior 
laterals 1.2. Anterior median eyes separat- 
ed by 1.4 diameters, posterior median eyes 
by 1.0, anterior median eyes separated 
from anterior laterals by 3.4 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.1 their diameters. Total length 7.8 
mm. Carapace 3.2 mm long, 2.8 wide. 
First femur 4.3 mm, patella and tibia 4.4, 
metatarsus 4, tarsus 1.2. Second patella 
and tibia 3.8 mm, third 2.1, fourth 3.2. 

Male holotype. Light reddish brown 
carapace, lighter around eyes and white 
mark in center (Fig. 96). Legs yellowish 
white, gradually growing darker toward 
distal ends of articles. Femur I with row 
of four macrosetae on anterior side; five on 
anteroventral side. Dorsal folium, venter, 
and sternum as in female (Figs. 96, 97). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 
1.2, posterior laterals 1.1. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 1.0, anterior median eyes 
separated from anterior laterals by 2.5 di- 
ameters of anterior 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 
2.2. 

Diagnosis. Unlike M. nigriventris or M. 



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 inm, 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. 

Distribution. Moderately high altitudes 
in northern Chile and southern Peru (Map 
7). 

Records Exmnined. 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°1'W, 11.x. 1955 (L. E. Peiia, 



42 Bulletin Museum of Coinparative Zoology, Vol. 157, No. 1 



IRSN); Antofagasta, 23°39'S, 70°24'W, 15.xi.l975 (L. 
E. Pena, AMNH); Quillagua, 21°39'S, 69°33'W, 
4.ii.l965 (L. E. Pena, MCZ), 21.i.l973 (W. C. Sedg- 
wick, MCZ); Rio Loa, 25 km S Quillagua, 21°54'S, 
69°33'W, 20.viii.l966 (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.iii.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.i.l973 (W. 
C. Sedgwick, MCZ); Chapiquilta, 19°18'S, 69°25'W, 
6.vi.l968 (L. E. Peiia, MCZ); Chiapa, 19°32'S, 
69°13'W, 24. iv. 1969 (L. E. Peiia, MCZ); Pica, 
20°30'S, 69°21'W, 26.i.l973 (W. C. Sedgwick, MCZ); 
Poroma, Quebrada de Tarapaca, 19°52'S, 69°11'W, 
20.iv.l969 (L. E. Peiia, MCZ); Quisama, 19°19'S, 
69°28'W, 5.vi.l968 (L. E. Peiia, MCZ). PERU Are- 
quipa: Arequipa, 16°23'56"S, 71°32'6"W, 15.vii.l955 
(A. Meza, MZSP); Chiguata, near Arequipa, 16°24'S, 
71°24'W, 15.ii.l948 (W. Weyrauch, MUSM). 

1 3. Metepeira calamuchita new species 
Figures 100-107, 308; IVIap 5 

Holotype. Male from Calamuchita, Cordoba, Argen- 
tina, 15.i.l955, J. M. Viana, in MACN. The specific 
name is a noun in apposition after tlie 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 
w^hite 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 calainuchita 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 Historij. 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 



Metepeira • Piel 43 



mostly lower altitudes east of the Andes 

(Map. 5). 

Records Examined. ARGENTINA Buenos Aires: 
Las Espaduiias [?], 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.l972 (Amarril- 
la, MACN). Cordoba: 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.l955 
(J. M. Viana, MACN); Mina Clareo, 31°43'S, 65°0'W, 
15.iv.l973 (Stiebel, MACN). Jtijuij: Iturbe, 22°59'S, 
65°21'W, 22.ii.1983 (L. E. Peila, 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.l967 (L. E. Pena, MCZ). 
Santiago del Estero: Santiago del Estero, 27°47'S, 
64°16'W (AMNH), 3.iv.l965'"(H. W. Levi, MCZ); 70 
km W Santiago, 27°47'S, 65°25'W, 3.iv.l965 (H. W. 
Levi, MCZ); Quebrachos: Sumampa, Parayacu [?], 
29°22'S, 63°28'W, 15.xi.l944 (Maldonado Bruzzone, 
MLP). 

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, 
Aisen province, Chile. Reddish brown car- 
apace with white setae, light around eyes, 
lateral posterior extensions, central arrow- 
shaped inark (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 browii with a white fleur-de- 
lis that reduces posteriorly (Fig. 113). Ven- 
ter brownish gray with wide median white 



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 iTiedians 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, Aisen province, 
Chile. Leg coloration as in feinale, except 
femur I mostly reddish browii with row of 
four macrosetae on anterior side; five on 
anteroventral side. Carapace, dorsum, ven- 
ter and sternum as in female (Figs. Ill, 
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 froixi 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 mixi, 
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 




-40 -35 

Latitude (Degrees North) 

Figure 109. Days on which mature M. galatheae were col- 
lected between 1937 and 1989 with latitude of the collection 
locality. Seasonality appears to be more restricted in southern 
regions than in northern regions. 

Scale of abscissa: -150 = August 3; 1 = January 1; 150 = 
May 30. 



scape with a corresponding variation in the 
position of the epigynal openings, it is 
nonetheless possible to distinguish the fe- 
males of M. galatheae from those of other 
species in the M. nigriventris group. 
Whether ventrally or posteriorly posi- 
tioned, the epigynal openings and the 
darkened shadows of the sclerotized re- 
ceptacles beneath them are relatively 
smaller than those of M. calamuchita, M. 
tarapaca, and M. nigriventris (compare 
Figs. 118-120 with Figs. 86, 94, 102). Also, 
M. galatheae lacks the swollen posterior 
lobes present in M. karkii (compare Figs. 
118-120 with Fig. 123). The shape of the 
embolus of M. galatheae varies significant- 
ly (compare Fig. 109 with Fig. 110). How- 
ever, unlike other species, the embolus of 
M. galatheae has a distinct, round and 
swollen protrusion (Fig. 117). 

Variation. Average body length of 80 fe- 
males examined 7.9 mm, range 4.8 to 12.5 
mm. Average body length of 16 males ex- 
amined 5.9 mm, range 3.5 to 8.1 mm. Epi- 
gyna vary considerably. Many, similar to 
the holotype, open ventrally and resemble 
a posteriorly widened version of M. tara- 
paca (Figs. 119, 120); cleared epigyna 
show relatively straight ducts connecting 
the epigynal openings with the seminal re- 
ceptacles. These have a short distance be- 
tween the openings and the hood of the 
scape. Others, usually in southern Chile, 
have posterior openings and look surpris- 



ingly different (Fig. 118); cleared epigyna 
show S -shaped ducts connecting the open- 
ings with the seminal receptacles. These 
have an extended wrinkled area between 
the openings and the hood of the scape. 
However, several females [e.g., CHILE 
Bio-Bio: Chilian, 8.xi.l976 (G. Moreno, 
AMNH); Las Lajuelas, ll.i.l976 (G. Mo- 
reno, AMNH)] have epigyna that appear 
to be intermediate between the two forms. 
Furthermore, no somatic features were 
found to be sufficiently different, and little 
corresponding variation was found among 
sympatric males. It is possible that further 
collecting efforts will discover correspond- 
ing males, and future molecular studies 
may show that speciation has, in fact, oc- 
curred. But in the meantime, I am opting 
to treat both varieties as belonging to the 
same species. 

Natural History. Although mature spec- 
imens have been collected throughout the 
year (Fig. 304), the seasonality of this spe- 
cies appears to depend on its latitude. At 
the 45th southern parallel, spiders are usu- 
ally found in late November and Decem- 
ber; at the 40th parallel, spiders occur be- 
tween October and Febrtiary; and at the 
30th parallel, they are collected year round 
(Fig. 108). Median elevation, 550 m. Spi- 
ders are found on Patagonian scrub, 
dunes, and wire fences. 

Distribution. Chile and Argentina (Map 
7). 

Records Examined. ARGENTINA Buenos Aires: 
Felipe Sola, 38°1'S, 62°50'W, 15.1.1944 (Prosen, 
MLP); Patagones, 40°48'S, 62°59'W, 15.ii.l937 (J. M. 
Viana, MACN); Sierra de la Ventana, 38°9'S, 
61°48'W, 15.iii.l939 (J. C. Gario, MACN). Catamar- 
ca: Mutquin, 28°19'S, 66°10'W, 15.1.1963 (O. de Fer- 
rariis, AMNH). Chubut: 15 km S Epuyen, 42°22'S, 
71°21'W, 15.1.1986 (P A. Goloboff, N. I. Platnlck, & 
R. T. Schuh, AMNH); 19.5 km E Shaman, 44°27'S, 
70°30'W, 19.xi.l966 (E. I. Schllnger & M. E. Irwin, 
CAS); 3 km N Puerto Lobos" 41°59'S, 65°6'W, 
14.xii.l966 (E. I. Schllnger & M. E. Irwin, CAS); 35 
km E Esquel, 42°54'S, 70°53'W, 18.xl.l966 (E. I. 
Schhnger & M. E. Irwin, CAS); El Hoyo [?], 42°4'S, 
71°30'W (A. Kovacs, AMNH), 10.1.1962 (Andor Ko- 
vacs, AMNH); Epuyen, 42°15'S, 71°23'W, 18.xl.l962 
(Andor Kovacs, AMNH); Leleque, 42°28'S, 71°6'W, 
12.11.1965 (Andor Kovacs, AMNH); Los Manantlales, 



Metepeira • Piel 



45 




A A 









Figures 110-120. Metepeira galatheae {ThoreW) (sp. 14 [110-114,116,117,120] 46°33'S, 71°57'W; [115,119] 29°50'S, 70°2'W; 
[118] 33°30'S, 71°25'W). 110, male palpus, mesal. Ill, male, dorsal. 112, male, ventral. 113, female, dorsal. 114, female, 
ventral. 115, epigynum, posterior. 116, epigynum, posterior. 117, male embolic division, ventral. 118-120, epigynum, ventral. 
Figures 121-128. Metepeira /car/f/V (Tullgren) (sp. 15; 51°38'S, 69°13'W). 121, male palpus, mesal. 122, epigynum, posterior. 
123, epigynum, ventral. 124, male, dorsal. 125, male, ventral. 126, female, dorsal. 127, female, ventral. 
Scale bars: dorsum and venter figures 1.0 mm. 



N of Comodoro-Rivadavia, 45°28'S, 69°29'W, 
19.xi.l985 (L. E. Peiia, AMNH); N of Camarones, 
Cantera, Namuncura, 44°46'S, 65°42'W, 17..>d.l9S5 
(L. E. Pena, AMNH); Rio Turbio, 42°13'S, 71°41'W 
(Andor Kovacs, AMNH), 12.1.1962 (Andor Kovacs, 
AMNH). Cordoba: 12 mi W Sampacho, 33°23'S, 
64°43'W, 7.ii.l951 (E. S. Ross & Michelbacher, CAS); 
Arguello, 31°21'S, 64°15'W, 15.xii.l943 (J. A. De Car- 



lo, MACN); Calamuchita, 32°4'S, 64°33'W, 
15.xii.l940 (J. M. Viana, MACN); Sampacho, 33°23'S, 
64°43'W, 7.ii.l951 (E. S. Ross & Michelbacher, CAS). 
Mendoza: Between Beazley and San Rafael, 34°10'S, 
67°29'W, 4.iii.l983 (L. E. Pefia, AMNH); Mendoza, 
32°53'S, 68°49'W, 30.iii.l965 (H. W. Levi, MCZ); Us- 
pallata, 32°35'S, 69°20'W, 7.iii.l983 (L. E. Pefia, 
AMNH). Neuquen: Catan Lil, Charaliuilla, 39°45'S, 



46 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



70°37'W, 15.ii.l971 (O. de Ferrariis, AMNH); Cuba 
del Leon [?], 39°9'S, 70°53'W, 15.1.1975 (Mauiy, 
MACN); Lago Alumine, 38°55'S, 71°9'W, 15.1.1976 
(O. de Ferrariis, AMNH); Zapala, 38°54'S, 70°4'W, 
15.1.1958 (J. R. Navas, MACN). Rio Negro: Cerro 
Alto [?], 41°8'S, 70°40'W (MACN); Co. Leones, 
source of Rio Limay, 40°33'S, 70°26'W, 28.11.1959 (J. 
R. Navas, MACN); El Bolson, 41°58'S, 71°31'W, 
1.11.1961 (A. Kovacs, AMNH), 17.X.1961 (Andor Ko- 
vacs, AMNH); El Bolson, 41°58'S, 71°35'W, 2.11.1965 
(Andor Kovacs, AMNH); Ceneral Roca, 39°3'S, 
67°32'W, 15.lx.1964 (Bachmann, MEG). Salta: Mauiy 
[?], 24°40'S, 65°45'W, 15.1.1975 (MACN). San Juan: 
10 km N Matagusanos, 31°10'S, 68°38'W, 13.1.1983 
(L. E. Peiia, AMNH). San Luis: INTA Experimental 
Station, E of Villa Mercedes, 33°40'S, 65°27'W, 
8.xil.l967 (C. R. Ward, CAS). Santa Cruz: 2.4 km S 
Fltz Roy, 47°2'S, 67°15'W, 12.xll.l966 (E. I. Schllnger 
& M. E. Irwin, CAS). Tucumdn: San Miguel de Tu- 
cuman, IML gardens, 26°49'S, 65°13'W, 19.xil.l979 
(L. A. Stange, FSCA). BRAZIL Mato Grosso: Campo 
Grande, 20°27'S, 54°37'W, 7.11.1952 (M. Alvarenga, 
MZSP). CHILE Aconcagua: W end tunnel, 85 km S 
Illapel, 32°49'S, 71°7'W, 29.xl.1950 (E. S. Ross & 
Mlchelbacher, CAS). Alsen: 8 km W Chile Chlco, 
46°33'S, 71°57'W, 22.xi.1966 (E. I. Schllnger & M. 
E. Ii^wln, CAS); Chile Chlco, near lake, 46°33'S, 
71°43'W, 21.xl.1966 (E. I. Schllnger & M. E. Invln, 
CAS). Antofagasta: 6 km N Muelle de Pledra, N Tal- 
tal, 25°21'S, 70°30'W, 4.11.1942 (Junius Bird, AMNH); 
Caleta Hueso Parado, Taltal, 25°22'S, 70°28'W, 
1.11.1941 (Junius Bird, AMNH); Coblja, 22°33'S, 
70°16'W (ZMUC); Quebrada Paposo, 25°2'S, 
70°27'W, 3.11.1989 (L. Stange, FSCA). Araucania: Pe- 
mehue [?], 38°3'S, 71°43'W (L. E. Pena, IRSNB); 
Villarrlca, 36°16'S, 72°13'W, 25.xl.1963 (G. F Ed- 
munds, AMNH). Atacama: 50-60 km S Copiapo, 
27°51'S, 70°20'W, 24.vlll.1966 (E. I. Schllnger & M. 
E. Irwin, CAS); Copiapo, 27°22'S, 70°20'W (Cartis, 
MNRJ); Rio Copiapo, by the sea, 27°19'S, 70°56'W, 
13.vl.1968 (L. E. Pena, MCZ). Bio-Bio: 4 km E road 
to Pinto, 36°42'S, 71°53'W, 4.1.1976 (B. Moreno, 
AMNH); Chilian, 36°36'S, 72°7'W, 2.1.1976 (G. Mo- 
reno, AMNH), 21.11.1978 (G. Moren, MCZ); Chilian, 
In cemetery, 36°36'S, 72°7'W, 8.xl.l976 (G. Moreno, 
AMNH); Cuesta de QuUmo, Chilian, 36°38'S, 
72°12'W, 13.xl.1976 (G. Moreno, AMNH); El Aban- 
Ico, 37°20'S, 71°31'W, 30.xll.l950 (E. S. Ross & 
Mlchelbacher, CAS); Las Lajuelas, 36°39'S, 72°8'W, 
11.1.1976 (G. Moreno, AMNH). Concepclon, 
36°50'S, 73°3'W (L. E. Pena, IRSNB); Concepclon: 
Salta de Rio Laja, 37°13'S, 72°23'W, 30.1.1951 (E. S. 
Ross & Mlchelbacher, CAS); Nuble: 50 km E San 
Carlos, 36°25'S, 71°6'W, 26.xll.1950 (E. S. Ross & 
Mlchelbacher, CAS); Nuble: Cordillera de Chilian 
[?], 36°51'S, 7r24'W, 1.11.1947 (L. E. Peiia, IRSNB); 
Rio Andallen, 36°44'S, 73°1'W, 25.111.1979 (S. Gu- 
tierrez, MCZ). Coquinibo: 20 ml E La Serena, 
29°54'S, 70°56'W, 3.vll.l950 (E. S. Ross & Mlchel- 
bacher, CAS); 5 ml N Ovalle, 30°31'S, 71°12'W, 
l.xil.l950 (E. S. Ross & Mlchelbacher, CAS); Banos 



del Toro, 29°50'S, 70°2'W, 15.11.1947 (L. E. Pena, 
IRSNB); Cerro Tahnay, 30°50'29"S, 71°37'14"W, 
29.xl.1961 (A. F. Archer, AMNH); Cuesta las Cardas, 
Ovalle Rd., 30°17'S, 71°16'W, 13.xl.l961 (R. Wagen- 
knecht, AMNH); Hacienda Illapel, 31°36'S, 71°7'W, 
3.X1.1954 (L. E. Pena, IRSNB), 19..X.1966 (E. I. 
SchUnger, M. E. Irwin, & L. E. Pena, CAS); Illapel: 
Salamanca: Fundo Quelen, 31°52'S, 70°52'W, 
30.lv.1961 (A. F. Archer, AMNH); La Serena, 
29°54'28"S, 7ri5'15"W, 15.11.1947 (L. E. Pena, 
IRSNB); Loma de Penuelas, 6 km S La Serena, 
29°57'S, 71°18'W, 28.xl.1961 (A. F. Archer, AMNH); 
Qullacan, 16 km E La Serena, 29°54'S, 71°5'W, 
2.x. 1961 (R. Wagenknecht, AMNH). Los Lagos: Purr- 
anque, 40°55'S, 73°10'W, 15.11.1955 (Edwin Reed, 
AMNH); Rio Bueno, 40°19'S, 72°58'W (L. E. Pena, 
IRSNB); Valdlvla: Neltume, 39°48'S, 71°57'W, 
23.xi.1988 (V. & B. Roth, CAS). Malleco: Angol, 
37°48'S, 72°43'W, 29.1.1951 (E. S. Ross & Mlchel- 
bacher, CAS). Maule: Linares, 35°51'S, 71°36'W (L. 
E. Pena, IRSNB), 15.1.1947 (L. E. Pena, IRSNB); 
Mlraflores, Pedag. [?], 35°55'S, 71°39'W (Toro, 
AMNH). O'Higgins: Cheplca, 34°44'S, 71°17'W, 
15..X11.1947 (L. E. Peiia, IRSNB). Region Metropoli- 
tana: 34 km W Sandago, 33°30'S, 71°25'W, 
19.xll.1950 (E. S. Ross & Mlchelbacher, CAS); Ba- 
tuco, nr. Santiago, 33°13'S, 70°47'W (Gull. Mann, 
AMNH); Lampa, 33°17'S, 70°54'W, l.v.1979 (L. E. 
Peiia, AMNH). Santiago: El Golf [?], 33°30'S, 
71°25'W, 9.iv.l961 (A. F Archer & J. Aros, AMNH); 
Santiago, 33°30'S, 71°25'W (L. E. Peiia, IRSNB), 
1.11.1973 (W. C. Sedgvdck, MCZ). Talca: 22 ml N Tal- 
ca, 35°7'S, 71°40'W, 22.xll.1950 (CAS). Valparaiso: 
Concon, in cow farm, 32°55'S, 71°31'W, 4.111.1962 (H. 
Morales, AMNH); La Cruz, 32°53'S, 71°16'W, 
18.1.1973 (W. C. Sedgwick, MCZ); Llay-Llay, 32°51'S, 
70°58'W, 20.1.1973 (W. C. Sedgwick, MCZ); Los Mai- 
tenes [?], 32°59'S, 71°15'W, 14.X.1954 (L. E. Peiiai 
IRSNB); Quintay, 33°11'S, 71°42'W, 19.11.1967 (E. I. 
Schhnger, CAS); Valparaiso, 33°2'S, 71°38'W (Edwin 
Reed, AMNH). 

15. Metepeira /car/c/7 (Tullgren) 
Figures 121-128, 303; Map 5 

Araneus karkii Tullgren, 1901: 219, 259. Female lio- 
lotype from Kark, Chile in the SMNH, examined. 

Metepeira labyrinthea: — Roewer, 1942: 868. Bonnet, 
1957: 2821. Erroneous synonymy. 

Description. Female from Rio Gallegos, 
Santa Cruz Province, Argentina. Carapace 
reddish brown with white setae, light 
around eyes with lateral posterior exten- 
sions (Fig. 127). Legs light yellow, articles 
annulated distally. Femur I with row of 
three to four macrosetae on anterior side; 
one to four on anteroventral side. Dorsum 
covered with denser, longer, black and 



Metepeira • Piel 



47 



white setae than in most species. Folium 
mostly white with brown speckles (Fig. 
127). Venter brownish gray with wide 
white median line; pair of large white spots 
on either side of spiracle. A pair of thin 
white lines, parallel to and on either side 
of median line, sometimes connect to a 
pair of thinner transverse white lines: one 
just posterior to the epigynal groove, one 
just anterior to the spinnerets (Fig. 128). 
Sternum dark reddish brown (Fig. 128). 
Ratio of eye diameters: posterior inedians 
and anterior medians 1.2, anterior laterals 
1.4, posterior laterals 1.5. Anterior median 
eyes separated by 1.8 diameters, posterior 
inedian eyes by 1.0, anterior median eyes 
separated from anterior laterals by 4.2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 9.5 inm. Carapace 4 mm long, 
3.4 wide. First femur 4 mm, patella and 
tibia 4.4, metatarsus 3.7, tarsus 1.2. Sec- 
ond patella and tibia 3.9 mm, third 2.5, 
fourth 3.8. 

Male from Rio Gallegos, Santa Cruz 
Province, Argentina. Carapace reddish 
brown with lighter eye region, lateral pos- 
terior extensions, and median arrowhead 
inark (Fig. 125). Legs light yellow, articles 
distally annulated reddish brown. Femur I 
with row of three to four macrosetae on 
anterior side; five to six on anteroventral 
side. Dorsal folium mostly white with 
brown speckles (Fig. 125). Venter and 
sternuiTi as in female (Fig. 126). Ratio of 
eye diameters: posterior medians and an- 
terior medians 1.0, anterior laterals 1.3, 
posterior laterals 1.2. Anterior inedian 
eyes separated by 1.7 diameters, posterior 
inedian eyes by 1.2, anterior median eyes 
separated from anterior laterals by 2.8 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.4 their diameters. To- 
tal length 5.7 mm. Carapace 2.8 mm long, 
2 wide. First femur 4.3 mm, patella and 
tibia 4.3, metatarsus 4.1, tarsus 1.2. Sec- 
ond patella and tibia 3.5 mm, third 1.9, 
fourth 2.9. 

Diagnosis. Females are easily separated 
from other species in the M. nigriventris 



group by the thick posterior epigynal lobes 
(compare Fig. 123 with Fig. 119). The dis- 
tal embolic apophysis does not protrude 
out from under the terminal apophysis 
(Fig. 124) as it does in M. galatheae, M. 
tarapaca, and M. nigriventris (Figs. 87, 95, 
119). The embolus of M. karkii differs 
from M. calaniuchita by lacking the in- 
wardly curved "dewlap" under the embo- 
lus (compare Fig. 121 with Fig. 103). 

Variation. Average body length of 13 fe- 
males examined 6.8 mm, range 5 to 8.2 
mm. Average body length of five males ex- 
amined 4.4 mm, range 2 to 5.3 mm. Dor- 
sal folia vaiy from white with little contrast 
and indistinct fleur-de-lis to darker with 
more contrast and distinct fleur-de-lis. 

Natural History. This species appears to 
be strongly seasonal: mature specimens 
have been collected between November 
and March (Fig. 303). Median elevation, 
300 m. Spiders are found in pampas (tree- 
less grassland). 

Distribution. Lower altitudes in south- 
ern Argentina and Chile (Map 5). 

Records Examined. ARGENTINA Chubut: Puerto 
Piramides, Peninsula Valdes, 42°34'S, 64°17'W, 
12.xi.l988 (V. & B. Rodi, CAS). Neuqtien: Laguna 
Blanca, 39°3'S, 70°23'W, 15.iii.l959 (J. Nara, 
MACN); Zapala, 38°54'S, 70°4'W, 15.1.1958 (J. R. Na- 
vas, MACN); Zapala, Laguna Blanca, 38°54'S, 
70°4'W, 15.1.1959 (J. R. Navas, MACN). Rto Negro: 
Cerro Alto [?], 41°8'S, 70°40'W (MACN); Coronel 
Juan Jose Comez, 39°2'S, 67°39'W, 15..\i.l945 (Ibarra 
Grasso, MLP); Ne-Luan, 41°25'S, 68°45'W (MACN). 
Santa Cruz: Laguna Calafate, Precordllfera [?], 
50°55'S, 70°9'W, 22.1.1967 (P San Martin, MCZ); Rio 
Gallegos, 51°38'S, 69°13'W, 20.1.1967 (P San Martin, 
MCZ). BOLIVIA Santa Cniz: Patagonia: Estancia 
Monte, cerca Rio Coyby [?], 50°14'S, 68°55'W (B. 
Brown, AMNH). CHILE Magallanes: 4 km W La- 
guna Amarga, 50°59'S, 72°49'W, 8.xil.l966 (E. I. 
Schlinger & M. E. Irwin, CAS); Kark, 51°17'30"S, 
72°32'30"W, 13.111.1899 (E. Nordenskiold, NRMS). 

Metepeira compsa Group 

The three species in the M. co77ipsa 
group include Metepeira compsa, Metepei- 
ra roraima, and Metepeira gressa. Along 
with the Metepeira nigriventris group, this 
group has a median apophysis with teeth 
on the face of the keel. Unlike the Mete- 



48 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



peira nigriventris group, this group has 
smaller, slimmer distal embolic apophyses 
that do not arch up over the embolus tip 
(compare Fig. 149 with Fig. 84). The 
openings to the epigynum have distinct, 
sclerotized edges and are clearly visible ei- 
ther as circles (Figs. 131, 134) or eye- 
shaped ovals (Figs. 143, 151). 

1 6. Metepeira compsa (Chamberlin) 
Figures 129-140, 311; Map 8 

Aranea compsa Chamberlin, 1916: 252, fig. 6, ? . Fe- 
male holotype from Ollantaitambo, Cusco, Peru, in 
the MCZ, examined. Bonnet, 1955: 462. 

Araneus lahijrintheus: — Petnmkevitch, 1926: 27. Er- 
roneous synonymy. 

Metepeira virginensis Chamberlin and Ivie, 1942: 70, 
figs. 188-190, 9,8. Female holotype from St. 
Thomas, U.S. Virgin Islands, in AMNH, examined. 
NEW SYNONYMY. 

Metepeira latigijna Chamberlin and Ivie, 1942: 70, 
figs. 191, 192, 5 . Female holotype from Porto Ale- 
gre [Baliia], Brazil, in AMNH, examined. NEW 
SYNONYMY. 

Metepeira compsa: — Chamberlin and Ivie, 1942: 71, 
figs. 193-195, $, c?. 

Metepeira labyrinthea: — Biyant, 1942: 346. 

Metepeira perezi Archer, 1965: 132, figs. 14, 16, 3, 
9 . Male holotype from subexperiment station, Is- 
abela, Puerto Rico, in AMNH, examined. Brignoli, 
1983: 276. NEW SYNONYMY. 

Metepeira vaurieonim Archer, 1965: 133, figs. 15, 19, 
S, ?. Male holotype from Usine de Robert, Mar- 
tinique, in AMNH, lost. BrignoU, 1983: 276. Lo- 
pez, 1993: 10, 11, figs. 1-4, 11, 6. NEW SYN- 
ONYMY. 

Note. Although the type for M. vaurieonim is 
lost, all examined records from Martinique (includ- 
ing ones from "Usine de Robert") belong to M. 
compsa. 

Description. Female from Savonet, Cu- 
rasao, Netherlands Antilles. Carapace light 
around eyes with lateral posterior exten- 
sions (Figs. 135, 137). Only tibia IV 
ringed. Femur I with row of four macro- 
setae on anterior side; two or three fine 
setae on anteroventral side. Dorsum of ab- 
domen with usual Metepeira folium, 
though whiter than usual in some speci- 
mens (Figs. 135, 137). Venter with wide 
white median line flanked by two thin 
(Fig. 136) or wide (Fig. 138) white lines; 
pair of white spots on either side of spi- 
racle. Sternum has wide median white line 



widening anteriorly, sometimes broken 
(Figs. 136, 138). Ratio of eye diameters: 
posterior medians and anterior medians 
1.0, anterior laterals 1.3, posterior laterals 
1.2. Anterior median eyes separated by 1.3 
diameters, posterior median eyes by 0.7, 
anterior median eyes separated from an- 
terior laterals by 1.8 diameters of anterior 
lateral eyes, lateral eyes separated by 0.2 
their diameters. Total length 3.9 mm. Car- 
apace 1.8 mm long, 1.4 wide. First femur 
1.9 mm, patella and tibia 1.9, metatarsus 
1.5, tarsus 0.7. Second patella and tibia 1.7 
inm, third 0.9, fourth 1.5. 

Male from Savonet, Curasao, Nether- 
lands Antilles. Carapace light around eyes 
with lateral posterior extensions. Slightly 
lighter median triangular mark anterior to 
thoracic furrow (Fig. 139). Legs lightly 
ringed. Femur I with row of four macro- 
setae on anterior side, four on anteroven- 
tral side. Dorsum, venter, and sternum as 
in female (Figs. 139, 140). Ratio of eye 
diameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.3, posterior 
laterals 1.2. Anterior median eyes separat- 
ed by 1.3 diameters, posterior median eyes 
by 0.6, anterior median eyes separated 
from anterior laterals by 1.4 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.3 their diameters. Total length 3.6 
mm. Carapace 1.8 min long, 1.4 wide. 
First femur 2.7 mm, patella and tibia 2.8, 
metatarsus 2.4, tarsus 0.9. Second patella 
and tibia 2.3 mm, third 1.2, fourth 1.8. 

Diagnosis. Unlike other species, the 
openings to the epigynum of M. compsa 
are small and almost perfectly round and 
sclerotized around the riin (Fig. 131). In 
Peruvian and Argentinean populations the 
scape can be extremely wide, often entire- 
ly covering a ventral view of the openings 
(Fig. 134). The distal embolic apophysis 
on the male palp extends straight from its 
base and parallel to the embolus tip (Figs. 
129, 132), in contrast to those of other spe- 
cies in the M. compsa species group that 
lift away from the embolus tip (Figs. 141, 
149). 

Variation. Average body length of 22 fe- 



Metepeira • Piel 49 




roraima 
(17) 



Figures 129-140. Metepeira compsa (Chamberlin) (sp. 16 [129-131,135,136,139,140] 12°20'N, 69°7'W; [132] 4°30'S, 81°8'W; 

[133,134] 4°51'S, 80°46'W; [137,138] 17°5'N, 61°42'W). 129, male palpus, mesal. 130, epigynum, posterior. 131, epigynum, 

ventral. 132, male palpus, mesal. 133, epigynum, posterior. 134, epigynum, ventral. 135, female, dorsal. 136, female, ventral. 

137, female, dorsal. 138, female, ventral. 139, male, dorsal. 140, male, ventral. 

Figures 141-148. Metepeira roraima new species (sp. 17 [141-147] 3°22'N, 60°19'W; [148]3°21'N, 76°33'W). 141, male palpus, 

mesal. 142, epigynum, posterior. 143, epigynum, ventral. 144, male, dorsal. 145, male, ventral. 146, female, dorsal. 147, female, 

ventral. 148, female, dorsal. 

Scale bars: dorsum and venter figures 1 .0 mm. 



males examined 5 mm, range 3.5 to 7.3 
mm. Average body length of 18 males ex- 
amined 3.6 mm, range 2.4 to 4.7 mm. 
Enormous variation in the shape of the 
scape can be seen in this species: popula- 
tions in the Caribbean and northeastern 
South America have a narrow scape (Fig. 
131), whereas populations in Argentina 



and Peru have a thick scape (Fig. 134). 
Somewhat less consistently parallel differ- 
ences can be seen in the shape of the me- 
dian apophysis: flagella are centered in Ca- 
ribbean and northeastern South America 
(Fig. 129) but shifted to the left in Argen- 
tina and Peru (Fig. 132). Further subtle 
differences appear in the shape of the em- 



50 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



bolus tip and embolus arm (compare Fig. 
129 with Fig. 132). These male and female 
genital differences are not sufficiently con- 
sistent to provide solid evidence for spe- 
ciation; thus, these populations will be 
treated as one species. 

Natural History. Mature speciinens 
have been collected throughout the year 
(Fig. 311). Altitudes range from just above 
sea level (for Caribbean and eastern South 
American localities) to 4,000 m (for Pe- 
ruvian localities). Spiders have been found 
in everything from second growth mixed 
exotics (e.g., mango, citrus, banana) and 
mangroves to grasses and semidesert chap- 
arral. 

Distribution. Puerto Rico to northern 
Argentina, but absent from the Amazon 
(Map 8). 

Records Examined. ARGENTINA Buenos Aires: 
Punta Lara, 34°49'S, 57°59'W, 15.ii.l941 (F. Moneos, 
MACN). Chaco: Basail, 27°52'S, 59°18'W, 18.iv.l942 
(MACN). Corrientes: Paso de la Patria, 27°19'S, 
58°35'W, 15.1.1966 (M. E. Galiano, MEG). Entre 
Rws: Parana, 31°44'S, 60°32'W (Rosenzwaig, MLP); 
Salto Grande, 31°13'S, 57°56'W, 15.iii.l964 (M. E. 
Galiano, MEG). Misiones: Posadas, 27°23'S, 55°53'W, 
15.ix.l963 (M. E. Galiano, MEG). Neuquen: Piedra 
del Aguila, 40°3'S, 70°5'W, 18.vii.l978 (Mision Cien- 
tifica Danesa, ZMUC). Santa Fe: Tostado, 29°14'S, 
61°46'W (A. Giai, MACN). BOLIVIA La Paz: Apolo, 
14°43'S, 68°31'W, 10.viii.l989 (L. E. Pefia, AMNH). 
BRAZIL Bahia: Arquipelago dos Abrolhos, 17°40'S, 
38°50'W, 28.xii.1887 (U.S. E.G., Voy. of Albatross, 
USNM); Parque Ondina, Salvador, 12°59'S, 38°31'W, 
25.vii.1962 (A. F. Archer, AMNH); Porto Alegre, 
18°5'S, 39°34'W (AMNH). Minas Gerais: Pedra /lzuI, 
16°1'S, 41°16'W, 15.xii.l970 (F. M. Oliveira, AMNH). 
Periiambuco: Pernambuco [?], 8°3'S, 34°54'W, 
12.iii.l927 (SMF), 8.iii.l955 (SMF). Rio de Janeiro: 
Lagomar [?], Macae, near sea, 22°23'S, 41°47'W, 
17.vii.l986 (R. L. C. Baptista, MZSP). Rio Grande do 
Sal: Montenegro, 29°42'S, 51°28'W, 3.xi.l977 (E. H. 
Buckup, MGN); Rambo [?], 30°4'0"S, 51°11'W 
(MNRJ); Sao Leopoldo, 29°46'S, 51°9'W, 14.X.1965 
(Gelia Valle, MZSP). Sao Paulo: Rubiao, Jr. [?], Bo- 
tucatu, 22°52'S, 48°26'W, 25.iv.1988 (R. L. G. Bap- 
tista, MZSP); Sao Paulo, Guamja, 24°0'S, 46°16'W, 
24.vii.1983 (R. Sievers, AMNH). BRITISH WEST 
INDIES Anegada: 18°45'N, 64°20'W, 12.xi.l966 
(Harry Beatty, AMNH); center, nr. salt pond, 
18°45'N, 64°20'W, 4.vi.l966 (Island Project Staff, 
Univ of Puerto Rico, AMNH); West end, 18°45'N, 
64°22'W 4.vi.l966 (Island Project Staff, Univ. of 
Puerto Rico, AMNH); Anegada Settlement: 18°45'N, 
64°20'W, 5.vi.l966 (Island Project Staff, Univ. of 



Puerto Rico, AMNH). Antigua: Goolidge Airport, 
17°6'N, 61°51'W, 15.xi.l967 (N. L. H. Krauss, 
AMNH); Devils Bridge, 17°5'N, 61°42'W, 30.vi.l963 
(Rick & E. N. Kjellesvig-Waering, AMNH); Jolly 
Beach, 17°4'N, 61°53'W 20.ix.l963 (E. N. Kjellesvig- 
Waering, AMNH); Lignum Vitae Bay: Jolly Beach [?], 
17°4'N, 61°53'W, 19.vi.l968 (E. N. Kjellesvig-Waer- 
ing, AMNH); Redonda Island, from webs spun be- 
tween boulders on beach, 16°55'N, 62°19'W, 
10.ivl9.56 (J. F G. Glarke, USNM); Reeds Point, nr. 
Jolly Beach; 17°4'N, 61°53'W, 2.vii.l963 (E. N. Kjel- 
lesvig-Waering, AMNH); Saint John's, 17°6'N, 
61°51'W, 15.viii.l967 (N. L. H. Krauss, AMNH), 
15.xii.l967 (N. L. H. Krauss, AMNH). Dead Mans 
Ghest [?]: 18°22'N, 64°.34'W, 26.vl966 (Island Pro- 
ject Staff, Univ. of Puerto Rico, AMNH). East Seal 
Dog: 18°30'N, 64°25'W 7.vi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). George Dog Is- 
land: 18°30'N, 64°27'W, 7.vi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH); 4 mi W Virgin 
Gorda Isl, high on diy cliffs, 18°30'N, 64°27'W, 
25.iii.1979 (K. Johnson, AMNH). Ginger Island: 
18°24'N, 64°28'W, 25.V.1966 (Island Project Staff, 
Univ of Puerto Rico, AMNH). Great Dog: 18°29'N, 
64°27'W, 7.vi.l966 (Island Project Staff, Univ. of 
Puerto Rico, AMNH). Green Cay, near Tortola: 
18°27'N, 64°42'30'W, 14.viii.l965 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Island O. near 
Anegada Setdement: 18°45'N, 64°20'W, 5.vi.l966 (Is- 
land'Project Staff, Univ. of Puerto Rico, AMNH). Is- 
land R. near Anegada Settlement: 18°45'N, 64°20'W, 
4.vi.l966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). ]ost Van Dyke: 18°28'N, 64°45'W, 
30.viii.l965 (H. Heatwole, R. Levins & F Mac- 
Kenzie, AMNH). Large mangrove patch nr. Settle- 
ment Anegado: 18°45'N, 64°20'W [?], 5.vi.l966 (Is- 
land Project Staff, Univ. of Puerto Rico, AMNH). Lit-\ 
tie Camanoe: 18°28'N, 64°33'W, 25.vl966 (Island 
Project Staff, Univ. of Puerto Rico, AMNH). Little 
Josi Van Dyke: 18°27'N, 64°43'W, 27.vii.1965 (Island 
Project Staff, Univ of Puerto Rico, AMNH). Little 
Tobago: 18°26'N, 64°51'W 4.iv.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Montserrat: 
Gage's Soufriere [?], 16°43'N, 62°10'W 28.vii.1972 
(N. L. H. Krauss, AMNH); Plymouth, 16°42'N, 
62°13'W, 15.xi.l967 (N. L. H. Krauss, AMNH). Neck- 
er Island: 18°33'N, 64°21'W, 6.vi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). nr. Anegada Set- 
tlement: Byer's Bache [?], 18°45'N, 64°20'W, 
5.vi.l966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). Peter Island: 18°22'N, 64°35'W, 6.vii.l965 
(Island Project Staff, Univ of Puerto Rico, AMNH), 
12.V.1966 (Percy Chubb, AMNH). Prickly Pear Is- 
land: 17°10'N, 61°48'W, 6.vi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Saint Christo- 
pher Island: Basseterre, 17°18'N, 62°43'W, 6.ii.l968 
(B. Malkin, AMNH). Salt Island: 18°23'N, 64°31'W, 
24.V.1966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). Sandy Key: near Tortola, 18°13'N, 63°7'W, 
31.viii.l965 (H. & A. Heatwole, AMNH). Tobago Is- 
land: 18°27'N, 64°48'W, 2.iv.l966 (Island Project 



Metepeira • Piel 



51 



Staff, Univ. of Puerto Rico, AMNH). Tortola: 
18°27'N, 64°36'VV, 8.vii.l958 (A. F. Archer, C. Hel- 
sley, & M. Sanderson, AMNH); Beef Island, 18°27'N, 
64°31'W, 21.vii.l965 (H. Heatwole, R. Levins & F. 
MacKenzie, AMNH); Greater Camanoe Isl, 18°29'N, 
64°32'W, l.vii.l965 (Island Project Staff, Univ. of 
Puerto Rico, AMNH); Little Thatch Island, 18°23'N, 
64°42'W, 16.viii.l965 (Island Project Staff, Univ. of 
Puerto Rico, AMNH); Long Bay Estate, 18°24'N, 
64°41'W, 24.vii.1965 (Island Project Staff, Univ. of 
Puerto Rico, AMNH); Marina Key [?], 18°27'N, 
64°36'W, 4.vii.l965 (H. Heatwole & R. Levins, 
AMNH); Prospect Reef, S Roadtown, 18°25'N, 
64°36'W, 23.iii.1979 (K. Johnson, AMNH), 31.iii.l979 
(K. Johnson, AMNH); Road to town, 18°27'N, 
64°36'W, 15.vii.l972 (N. L. H. Krauss, AMNPI); 
Sandy Spit [?], 18°27'N, 64°36'W, 14.viii.l965 (Island 
Project Staff, Univ. of Puerto Rico, AMNH). Toi-tola 
Island: 18°27'N, 64°36'W, 15.vii.l965 (Island Project 
Staff, Univ of Puerto Rico, AMNH). Virgin Gorda: 
18°30'N, 64°24'W, 15..xi.l966 (Hany Beatty, AMNH); 
Baths & Devils Bay, 18°26'N, 64°27'W, 25.vi.1966 
(Island Project Staff, Univ. of Puerto Rico, AMNH); 
Cooper Mine Trail 18°26'N, 64°26'W, 25.vi.1966 (Is- 
land Project Staff, Univ. of Puerto Rico, AMNH); Sa- 
vana Bay & Pond Bay, 18°28'N, 64°25'W, 27.vi.1966 
(Island Project Staff, Univ. of Puerto Rico, AMNH); 
Trelhs Bay (W end of island), nr. harbor, 18°27'N, 
64°26'W, 27.iii.1979 (K. Johnson, AMNH); Virgin 
Gorda Mountain, 18°30'N, 64°24'W, 26.vi.1966 (Is- 
land Project Staff, Univ. of Puerto Rico. AMNH). 
Virgin Islands Area: Island Q [?], 18°45'N, 64°20'W, 
5.vi.l966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). West Seal 'Dog Island: 18°29'N, 64°28'W, 
7.vi.l966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). CHILE Tarapaca: Arica, 18°29'S, 70°20'W, 
28.i.l973 (W. C. Sedgwick, MCZ); Azapa Arica, 
18°31'S, 70°11'W, 9..xi.l955 (L. E. Peiia, IRSNB); 
Lluta, 18°24'S, 70°19'W, 12.xi.l955 (L. E. Pena, 
IRSNB); Rio Lluta, 18°24'S, 70°19'W, I2.xi.l955 (L. 
E. Pena, IRSNB); Sobraya, 18°32'S, 70°9'W, 
10.xi.l955 (L. E. Peila, IRSNB); Taltape, Camarones 
Valley, 18°59'S, 69°47'W, 29.i.l973 (MCZ). COLOM- 
BIA Magdalena: Cabaila "Villa Culebra," 10 km E 
Station Marta, 11°12'N, 74°7'W, 15.X.1985 (H.-G. 
Miiller, SMF); Casajera [?], 11°0'N, 74°15'W, 
l.ii.l974 (J. A. Kochalka, JAK); Cienaga, 11°1'N, 
74°15'W, 30.i.l974 (J. A. Kochalka, JAK); Gaira, 
11°11'N, 74°13'W, 15.xii.l975 (W. Eberhard, MCZ). 
DOMINICAN REPUBLIC Barahona: Patos, 
10°38'N, 61°52'W, 24.ix.1944 (R. H. Montgomery, 
AMNH). FRENCH WEST INDIES Guadeloupe: 
Deshaies, 16°18'N, 61°48'W, 28.vi.1960 (C. & P Vau- 
rie, AMNH); Domaine Duclos [?], 16°16'N, 61°31'W, 
25.vi.1960 (C. & P Vaurie, AMNH), 15.vii.l960 (C. 
& P. Vaurie, AMNH); Marie-Galante, in citrus, 
15°56'N, 61°16'W, 15.iii.l977 (W. H. Whitcomb, 
FSCA); Pointe-a-Pitre, Ilet a Boissard, 16°14'N, 
61°34'W, 26.vi.1960 (C. & R Vaurie, AMNH); Terre- 
de-Haut, Les Saintes, 15°58'N, 61°35'W, 2.vii.l960 
(C. & P. Vaurie, AMNH). Martinique: Ansemitian [?], 



Trois-Ilets, 14°33'N, 61°2'W, 10.vi.l960 (C. & P Vau- 
rie, AMNH); Diamant, 14°29'N, 61°2'W, 17.vi.l960 
(C. & P Vaurie, AMNH), 18.vi.l966 (C. & P Vaurie, 
AMNH); Fort de France, 14°36'N, 61°5'W, 
15.xii.l950 (N. L. H. Krauss, MCZ); Pointe Ferret, 
La Caravelle, 14°45'N, 60°54'W, 19.vi.l960 (C. & R 
Vaurie, AMNH); Sainte-Anne, 14°26'N, 60°53'W, 
20.vi.l966 (C. & P Vaurie, AMNH); Usine de Robert 
[?], 14°41'N, 60°57'W, 16.vi.l960 (C. & R Vaurie, 
AMNH). GRENADA nr Saint Georges: 12°3'N, 
61°45'W, 3.vi.l950 (Leo Isaacs, AMNH). NETH- 
ERLAND ANTILLES Bonaire: Red Pond [?], 
12°12'N, 68°15'W, 3.i.l968 (B. Malkin, AMNH). Cu- 
ragao: 12°11'N, 68°58'W, 13. i. 1968 (B. Malkin, 
AMNH); Fuik (Oostpunt), muddauber nests, 12°4'N, 
68°49'W, 20.xii.l962 (H. W. Levi & B. de Jong, 
MCZ); Groot Sint Joris, plantation, 12°14'N, 69°3'W, 
22.xii.1962 (H. W. Levi & B. de Jong, MCZ); Groote 
Berg, 12°11'N, 69°0'W, 19.xii.l962 (H. W. & L. Levi, 
MCZ); Hato, 12°11'N, 68°58'W, 28.xii.1967 (B. Mal- 
kin, AMNH), 7.i.l968 (B. Malkin, AMNH); Piscadera 
Baai, 12°8'N, 68°59'W, 18.xii.l962 (H. W. Levi, 
MCZ), 20.xii.l962 (H. W. Levi, MCZ); Savonet; 
shady ravine, 12°20'N, 69°7'W, 28.xii.1962 (H. W. 
Levi, MCZ); SE of airport, 12°10'N, 68°54'W, 
20.xii.l962 (H. W. Levi & B. de Jong, MCZ); Siberie, 
12°14'N, 69°3'W, 25.xii.1962 (H. W. Levi, MCZ); Sint 
Jan, 12°15'N, 69°6'W, 25.xii.1962 (H. W. Levi & B. 
de Jong, MCZ); Willemstad, Jewish Cemetary, 
12°7'N, 68°57'W, 24.xii.I962 (H. W. Levi, MCZ). Sint 
Eustatius: Oranjestad, 17°29'N, 62°59'W, 18.i.l968 
(B. Malkin, AMNH). Sint Maarten: nr. Juliana Air- 
port [?], 18°4'N, 63°4'W, 24.ii.1965 (H. Heatwole & 
F. MacKenzie, AMNH). PARAGUAY Alto Parana: 
Taguarazaya [?], 25°30'S, 54°50'W (AMNH). PERU 
Ancash: Huaraz, 9°32'S, 77°32'W, 6.xii.l980 (C. Gold, 
CAS). Apuriniac: 35 mi E Abancay, 13°38'S, 72°22'W, 
5.iii.l951 (E. S. Ross & Michelbacher, CAS); 40 mi 
E Abancay, 13°38'S, 72°20'W, 4.iii.l951 (E. S. Ross 
& Michelbacher, CAS); Abancay, 13°38'2"S, 
72°52'53"W, 6.iii.l951 (E. S. Ross &' Michelbacher, 
CAS). Cajamarca: Jaen, 5°42'34"S, 78°48'32"W, 
17.V.1967 (A. F. Archer, AMNH). Cusco: 40 mi W 
Cusco, 13°32'S, 72°33'W, 5.iii.l951 (E. S. Ross & 
Michelbacher, CAS); Hacienda Urco, near Galea, 
13°22'S, 71°54'W, 19. ix. 1939 (Karl P Schmidt, 
AMNH); Huacerpay [?], 13°37'S, 72°13'W, 10.ix.l993 
(J. Ochoa Camara, MCZ); Ollantaitambo, 13°15'17"S, 
72°15'48"W, 15.vii.l911 (Yale Pemvian Expedition, 
AMNH), 15.xii.l980 (C. Gold, CAS); Pisac, 
13°25'21"S, 71°50'48'W, 13.-xii.l980 (C. Gold, CAS); 
Urubamba, 13°18'28"S, 72°6'55"W, 15.i.l965 (Carras- 
co, MCZ), 18.ii.l965 (H. W. Levi, MCZ), 6.viii.l987 
(D. Silva, MUSM). Huanuco: Huanuco, 9°55'S, 
76°14'W, 6.X.1946 (J. C. Palhster, AMNH). lea: km 
367 between lea and Nazca, sandy semidesert, 
14°24'S, 75°23'W, 22.i.l952 (W. Weyrauch, CAS); 
Nazca, 14°50'S, 74°57'W, 15. iv. 1951 (P. Aguilar, 
GAS). Piura: Gerro Prieto, La Brea, 4°41'S, 81°6'W 
(CAS); Mallares, 4°51'S, 80°46'W, 8.vi.l941 (H. E. F. 
& D. E. F, CAS), 13.vii.l941 (H. E. F. & D. E. F, 



52 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



CAS); N of Negritos, Farinas Valley, 4°41'S, 81°18'W, 
9.X.1938 (D. L. & H. E. Frizzell, CAS); nr. Negritos, 
4°42'S, 81°18'W, 5.m.l939 (H. E. F, CAS); nr. Se- 
chura, 5°33'S, 80°51'W, 4.xi.l941 (D. L. F, CAS); Ne- 
gritos, 4°42'S, 81°18'W, 15.xi.l934 (H. E. F, CAS), 
15.iii.l941 (H. E. F, CAS); Farinas, 4°30'S, 81°8'W, 
7.V.1939 (D. L. & H. E. Frizzell, CAS), 31.V.1939 (D. 
L. & H. E. Frizzell, CAS), 15.X.1939 (D. L. & H. E. 
Frizzell, CAS); Farinas Valley, 4°30'S, 81°8'W, 
6.iii.l939 (D. L. & H. E. Frizzell, CAS), 8.iv.l939 (D. 
L. & H. E. Frizzell, CAS); Fortachuelo, 4°59'S, 
79°54'W, 29.iv.1939 (M. H. & H. E. Frizzell, CAS); 
Quebrada Mogollon, 4°32'S, 81°4'W, 30.iv.l939 (D. 
L. & H. E. Frizzell, CAS), ll.vi.l939 (D. L. & H. E. 
Frizzell, CAS); Rio Chira Valley, N of Amotape, 
4°53'S, 81°1'W, 13,xi.l938 (D. L. & H. E. Frizzell, 
CAS); S of Sechura, 5°33'S, 80°51'W, 25.X.1941 (D. 
L. F, CAS); San Lorenzo [?], Zona Alta Este Herrera, 
5°4'S, 79°47'W, 12.iv.l969 (F Aguilar, MCZ); Sullana, 
4°53'S, 80°41'W, 8.X.1939 (D. L. & H. E. Frizzell, 
CAS), 5.X.1941 (D. L. & H. E. Frizzell, CAS). Tum- 
bes: 34 km E, 25 km N Funta Farinas, 4°18'S, 
80°51'W, l.i.l939 (D. L. & H. E. Frizzell, CAS). 
SAINT LUCIA Castries: 14°1'N, 61°0'W, 28.vii.1963 
(E. N. Kjellesvig-Waering, AMNH). TRINIDAD & 
TORAGO Tohago Island: Ruccoo Ray, 11°10'N, 
60°48'W, 15.viii.l965 (E. N. Kjellesvig-Waering, 
AMNH); Guayaguayare Foint, 10°8'N, 61°2'W, 
14.ix.l963 (E. N. Kjellesvig-Waering, AMNH); Pi- 
geon Foint, 11°10'N, 60°50'W, 18.viii.l937 (E. D., 
MCZ); Salybia Ray, by shore, 10°42'N, 61°2'W, 
15.ii.l972 (J. A. L. Cooke, MCZ); Toco, 10°50'N, 
60°57'W, 19. iv. 1964 (Erik N. Kjellesvig-Waering, 
AMNH). Trinidad: Gasparee, 10°46'N, 61°19'W, 
3.xi.l944 (R. Montgomery, AMNH). URUGUAY Co- 
lonia: Fmita Gorda [?], 34°28'S, 57°51'W, 25.ii.1968 
(R. Capocasale & L. Rruno, CAS). USA Puerto Rico: 
Algodones Key, 18°12'N, 65°41'W, 15.x. 1964 (H. 
Heatwole, R. Levins & F MacKenzie, AMNH); Ral- 
neario [?] Guajataca, 18°21'N, 66°55'W, 4.vii.l958 (A. 
F Archer, AMNH); Railos de Coamo, 17°59'N, 
67°3'W, 2.iv.l990 (H. W & L. Levi, MCZ), 3.iv.l990 
(H. W & L. Levi, MCZ); Rarranquitas, 18°11'N, 
66°18'W, 28.xii.1977 (J. Coddington, USNM); below 
Quebradillas along old RR track, 18°28'N, 66°56'W, 
30.iii.l989 (H. W & L. Levi, MCZ); Cabeza de Ferro 
Island, 18°15'N, 65°35'W, 16.i.l965 (H. Heatv^^ole & 
F. MacKenzie, AMNH); Cayo Don Luis, 17°57'N, 
66°58'W 12.i.l966 (Island Froject Staff, Univ. of 
Fuerto Rico, AMNH); Cayo Norte, off Culebra, 
18°20'N, 65°15'W, 14.ivl965 (H. Heatwole & F. 
MacKenzie, AMNH); Cayo Pinerito, 18°15'N, 
65°36'W, 24.ix.1964 (H. Heatwole & F. MacKenzie, 
AMNH); Channel at Culebra, Isla del Diablo, 
18°23'N, 65°40'W, 12.viii.l965 (Island Froject Staff, 
Univ. of Fuerto Rico, AMNH); Cuevas de los Alfaros, 
Rarrio Mora, 18°29'N, 67°1'W, 20.vii.l958 (A. F. Ar- 
cher & Rolle, AMNH); Culebra Island, 18°19'N, 
65°17'W, 19.vii.l965 (F. MacKenzie, AMNH); Cule- 
bra, near Dewey, 18°18'N, 65°18'W, 10.viii.l965 (Is- 
land Froject Staff, Univ. of Fuerto Rico, AMNH); Cu- 



lebrita Island, 18°19'N, 65°14'W, 15.ivl965 (H. Hea- 
twole & F MacKenzie, AMNH), ll.viii.l965 (Island 
Froject Staff, Univ. of Fuerto Rico, AMNH); Dese- 
cheo Is, 18°23'N, 67°29'W, 19.ii.l914 (AMNH); De- 
secheo Island, 18°23'N, 67°29'W, 28.V.1965 (H. Hea- 
twole, R. Levins & F MacKenzie, AMNH); Faro de 
Cabo Rojo, 18°5'N, 67°9'W, 13.iii.l961 (F. Rolle, 
AMNH); Frank Key, Isl. #13 nr. La Farguera area, 
17°58'N, 67°3'W, 14.i.l966 (Island Froject Staff, 
Univ. of Fuerto Rico, MCZ); Heatwole Island, off 
Culebrita, 18°19'N, 65°13'W, 14.iv.l965 (H. Heatwo- 
le & F. MacKenzie, AMNH); Hormiqueros, 18°9'N, 
67°8'W, 16.ii.l962 (Aida Velez, AMNH), ll.iii.l962 
(Aida Velez, AMNH), 19.iii.l962 (Aida Velez, 
AMNH); Isabela, subexperiment station, 18°30'N, 
67°1'W, 17.viii.l957 (A. F. Archer, AMNH); Jayuya, 
coffee plantation, 18°13'N, 66°37'W, 23.iii.1986 (H. 
W. & L. Levi, MCZ); Juana Diaz, 18°3'N, 66°31'W, 
7.xi.l971 (J. E. Carico, JEC); Levins Rock [?], 
18°12'N, 65°41'W, 15.X.1964 (H. Heatwole, R. Levins 
& F. MacKenzie, AMNH); Loma Tinaja [?], S of La- 
guna Cartagena, 17°59'N, 67°6'W 5.vii.l958 (A. F. 
Archer & M. Sanderson, AMNH); Luquillo, near 
beach, 18°23'N, 65°44'W, 24.i.l932 (A. S. Mills, 
AMNH), 24.xii.1985 (V. & R. Roth, CAS), 27.iii.1988 
(H. W & L. Levi, MCZ); Mayagiiez, 18°12'N, 
67°9'W, 15.vii.l958 (A. F. Archer, MCZ); Mayagiiez, 
5 km N university campus, 18°12'N, 67°9'W, 5.i.l964 
(MCZ); Mayagiiez: Las Mesas, 18°11'N, 67°6'W, 
20.xi.l960 (F. Rolle, AMNH); Mayagiiez, university 
farm N university campus, 18°12'N, 67°9'W, 7.ii.l964 
(MCZ); McKenzie Key, Key #3 [?], 18°12'N, 
65°41'W, 29.X.1964 (H. Heatwole, R. Levins & F. 
MacKenzie, AMNH); Mona Island, Serralles, 18°5'N, 
67°54'W [?], 7.ivl944 (Reatty, MCZ); Muertos Is- 
land, beating bushes, 17°54'N, 66°32'W 28.vl959 
(Jordan & Martorell, AMNH); Muertos Island, mud) 
nests of Sceliphron caementarium, 17°54'N, 66°32'W, 
27.V.1959 (Medina & Martorell, AMNH); N slope Ti- 
naja, nr. Cartagena Lagoon, beating and sweeping, 
18°23'N, 67°10'W, 5.vii.l958 (M. W Sanderson, 
AMNH); Palominitos, 18°20'N, 65°34'W, 16.vi.l965 
(Island Froject Staff, Univ. of Fuerto Rico, AMNH); 
Falomino Island, 18°21'N, 65°34'W, 7.xi.l964 (H. 
Heatwole & F. MacKenzie, AMNH); Farguera, 
17°59'N, 67°3'W, 25.iii.1990 (H. W & L. Levi, MCZ); 
Fatillas, 18°0'N, 66°1'W, 3.iv.l931 (Mills & Leonard, 
AMNH); Fico Atalaya, Rte. 2, nr Aiiasco, 18°18'N, 
67°11'W, 3.vii.l958 (M. W. Sanderson, AMNH); Fi- 
fieros Island, 18°15'N, 65°35'W, 24.ix.1964 (H. Hea- 
twole & F. MacKenzie, AMNH); Flaya de Humacao, 
18°10'N, 65°45'W, 23.V.1964 (MCZ); Quebradillas, 
around hotel, 18°29'N, 66°56'W, 30.iii.l989 (H. W. & 
L. Levi, MCZ); Ratones Island, 17°56'N, 66°17'W, 
19.xi.l964 (H. Heatwole & R. Levins, AMNH); Rom- 
ero III [?], 17°57'N, 67°0'W, 13.i.l966 (Island Froject 
Staff, Univ. of Fuerto Rico, AMNH); Rubianes, 
Caimito Rajo [?], Cord. Jaicoa, 18°26'N, 67°8'W, 
19.vii.l958 (A. F Archer & Rolle, AMNH); Santurce, 
18°27'N, 66°4'W, 28.iii.1931 (A. S. Mills, AMNH); 
South of Corozo, Cabo Rojo salt flats, edge of salt 



Metepeira • Piel 



53 



ponds, 17°56'N, 67°11'W, 23.iii.1990 (H. W. & L. 
Levi, MCZ); Valle de Lajas, 18°1'N, 67°8'W, 3.vi.l958 
(A. F. Archer, AMNH); xerie hills N of Guanica, 
18°0'N, 66°55'W, 28.viii.1957 (A. F. Archer, AMNH); 
Zancudo Island (Isleta Marina), 18°20'N, 65°37'W, 
2..xi.l964 (H. Heatwole & F. MacKenzie, AMNH). 
U.S. VIRGIN ISLANDS Big Cockroach: 18°24'N, 
65°4'W, 7.vi.l966 (Island Project Staff, Univ. of 
Puerto Rico, AMNH). Congo Cay: 18°22'N, 64°48'W, 
12.xi.l966 (Island Project Staff, Univ. of Puerto Rico, 
AMNH). Grass Cay: 18°22'N, 64°50'W, 12.xi.l966 
(Univ. of Puerto Rico, AMNH). Great Saint James 
Island: 18°19'N, 64°50'W, 13.xi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Hans Lollik Is- 
land: 18°24'N, 64°55'W, 6.iv.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Little Cock- 
roaches: 18°25'N, 65°3'W, 7.vi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Little Hans Lol- 
lick: 18°25'N, 64°54'W, 5.iv.l966 (Island Project Staff, 
Univ. of Puerto Rico, AMNH). Little Saint James Is- 
land: 18°18'N, 64°50'W, 13.xi.l966 (Island Project 
Staff, Univ. of Puerto Rico, AMNH). Mingo Cay: 
18°22'N, 64°49'W, 12.xi.l966 (Island Project Staff, 
Univ. of Puerto Rico, AMNH). Saint John: 18°20'N, 
64°45'W, 9.iii.l925 (F. E. Lutz, AMNH), ll.vii.l958 
(A. F Archer & M. Sanderson, AMNH), 15.xii.l967 
(N. L. H. Krauss, AMNH); Cruz Ray, 18°20'N, 
64°48'W, 27.ii.1964 (A. M. Chickering, MCZ); nr. 
Cinnamon & Hart Bays on W half of island, 18°20'N, 
64°46'W, 2.vlii.l976 (D. E. & D. N, Rosen, AMNH). 
Saint Thomas: 18°24'N, 64°55'W, 24. ii. 1925 
(AMNH), 24.vii.1925 (A. Petrunkevitch, CAS), 
24.xi.1925 (AMNH); Crov^^n Mountain, 18°21'N, 
64°58'W, 30.viii.l957 (A. F. Archer, AMNH), 
7.vii.l958 (A. F. Archer, AMNH); Denmark Hill [?], 
18°24'N, 66°55'W, l.ix.l957 (A. F Archer, AMNH); 
Flagstok Hill, Stumpy Bay, 18°22'N, 65°0'W, 
9.vii.l958 (A. F. Archer, AMNH); Harmans, Char- 
lotte Amalia, 18°21'N, 64°56'W, 2.ix.l957 (A. F. Ar- 
cher & family, AMNH); Hassell Island, 18°20'N, 
64°56'W, l.ix.l957 (A. F Archer, AMNH), 15.ii.l964 
(A. M. Chickering, MCZ), 10.viii.l966 (M. L. Pres- 
sick, AMNH). St. Croix: Christiansted, 17°45'N, 
64°42'W, 3.vi.l911 (AMNH), 15.i.l955 (A. M. Nad- 
ler, AMNH); East End, 17°45'N, 64°40'W, 15.xii.l965 
(Island Project Staff, Univ. of Puerto Rico, AMNH); 
Green Key, 17°46'N, 64°40'W, 17.iv.l964 (H. Hea- 
twole, MCZ); Mount Eagle, dryish forest, 17°46'N, 
64°49'W, 15.xii.l965 (Island Project Staff, Univ. of 
Puerto Rico, AMNH); Protestant Cay, 17°45'N, 
64°42'W, 18.iv.l964 (H. Heatwole, MCZ); St. Croix, 
17°45'N, 64°54'W, l.ix.l966 (Chickering, MCZ), 
6.ix.l966 (Chickering, MCZ). VENEZUELA Depen- 
dencias Federales: Patos, 10°38'N, 61°52'W, 
23.ix.1944 (R. H. Montgomery, AMNH). Districto 
Federal: Punta Tanaguarena, in coastal bldg. & gar- 
den, 10°37'N, 66°48'W, 26.xii.1970 (W. B. Peck, 
CAS). 



1 7. Metepeira roraima new species 
Figures 141-148, 316; Map 8 

Holotype. Female from Rio Suioimu, Roraima, Brazil, 
X.1966, M. Abrorenga, in MZSP The specific name 
is a noun in apposition after the locality. 

Description. Female holotype. Brown 
carapace; lighter around eyes (Fig. 146). 
Coxae, femora, tibiae, and patellae tan, 
lighter ventrally Metatarsi, tarsi white. Fe- 
mur I with row of three or four macrose- 
tae on anterior side; none on anteroventral 
side. Dorsal folium with typical Metepeira 
fleur-de-lis pattern (Fig. 146). Venter of 
abdomen brown with wide white median 
line, flanked by thinner white lines that to- 
gether form a T-shape posteriorly (Fig. 
147). Pair of small white spots on either 
side of T-shape mark. Sternum brown with 
median white line (Fig. 147). Ratio of eye 
diameters: posterior medians and anterior 
medians 1.1, anterior laterals 1.5, posterior 
laterals 1.3. Anterior median eyes separat- 
ed by 1.2 diameters, posterior median eyes 
by 0.6, anterior median eyes separated 
from anterior laterals by 1.9 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.1 their diameters. Total length 4 mm. 
Carapace 1.8 mm long, 1.4 wide. First fe- 
mur 1.9 mm, patella and tibia 2.1, meta- 
tarsus 1.8, tarsus 0.7. Second patella and 
tibia 1.7 mm, third 1.0, fourth 1.6. 

Male paratype from Rio Surumu, Ro- 
raima, Brazil. Carapace brown; lighter 
around eyes with median white mark (Fig. 
144). Legs brown, white at base of femora. 
Femur I with row of four to five macro- 
setae on anterior side; four to five on an- 
teroventral side. Dorsal folium, venter, and 
sternum as in female (Figs. 144, 145). Ra- 
tio of eye diameters: posterior medians 
and anterior medians 1.1, anterior laterals 
1.6, posterior laterals 1.4. Anterior median 
eyes separated by 1.4 diameters, posterior 
median eyes by 0.6, anterior median eyes 
separated from anterior laterals by 1.6 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 2.9 mm. Carapace 1.3 mm long, 
1.1 wide. First femur 1.9 mm, patella and 



54 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



tibia 2, metatarsus 1.6, tarsus 0.6. Second 
patella and tibia 1.5 mm, third 0.8, fourth 
1.3. 

Diagnosis. The epigynum of M. roraima 
differs from that of M. compsa by having 
oval openings (Fig. 143) instead of round 
openings (Figs. 131, 134); it differs from 
M. gressa by being more translucent and 
by having a narrower scape (compare Fig. 
143 with Fig. 151). Unlike M. gres.sa, the 
embolus tip of M. roraima is more bent 
and the distal apophysis is as wide as the 
embolus tip (compare Fig. 141 with Fig. 
149). Unlike M. compsa, the distal embolic 
apophysis projects away from the embolus 
tip (Fig. 141) instead of extending straight 
from the base and parallel to the embolus 
tip (Figs. 129, 132). 

Variation. Average body length of eight 
females examined 5 mm, range 4 to 7 mm. 
Average body length of five males exam- 
ined 3.1 mm, range 2.8 to 4 mm. Colom- 
bian specimens are much darker than 
those from northern Brazil and southern 
Guyana, suggesting the possibility that 
these populations represent separate cryp- 
tic species. In fact, the carapace on Col- 
ombian specimens is often jet black with 
the white marks on the dorsal folium 
much reduced (Fig. 148). However, there 
is little corresponding genitalic difference, 
especially among males; consequently, 
these populations are deemed to be con- 
specific. 

Natural History. Mature M. roraima 
specimens from eastern South America 
have been collected in November and De- 
cember; specimens from western South 
America have been collected between 
March and July (Fig. 316). Median eleva- 
tion, 1,000 m. 

Distribution. Western Colombia, north- 
ern Brazil, and southern Guyana (Map 8). 

Records Examined. BRAZIL Roraima: Rio Suru- 
mu, Jerrit Rio Branco [?], 3°22'N, 60°19'W, x.1966 
(M. Abrorenga, MZSP). COLOMBIA. Valle del Cau- 
ca: Atuncela, 3°46'N, 76°42'W, 19.vii.l970 (W. Eber- 
hard, MCZ); Cali, around house, 3°27'N, 76°31'W 
(W. Eberhard, MCZ); Palmira, 3°32'N, 76°17'W, 
l.iii.l964 (Ballo, CAS); Rio Panee, below Buenos Ai- 



res, 3°55'N, 76°8'W, 5.iv.l970 (W. Eberhard, MCZ); 
Rio Pance, near Cali., 3°21'N, 76°33'W, 8.V.1970 (W. 
Eberhard, MCZ), 8.vi.l970 (W. Eberhard, MCZ), 
15.vi.l970 (W. Eberhard, MCZ), 23.vi.1970 (W. 
Eberhard, MCZ), 25.vi,1973 (W. Eberhard, MCZ). 
GUYANA Upper Takiitu: Isherton, on lat. 2, 10 mi E 
Rupununi River, 2°19'N, 59°22'W, 15.xi.l937 (W. G. 
Hassler, AMNH). 

18. Metepeira gressa (Keyserling) 
Figures 149-156, 300; Map 8 

Epeira gressa Keyserling, 1892: 166, fig. 123, 9 . Five 
female syntypes from Taquara, Rio Grande do Sul, 
Brazil, in BMNH, examined. One specimen des- 
ignated lectotype. 

Epeira seditiosa Keyserling, 1893: 212, fig. 157, S. 
Male holotype from Rio Grande do Sul, Brazil, in 
BMNH, examined. NEW SYNONYMY. 

Araneus gressus: — Petrunkevitch, 1911: 314. Roewer, 
1942: 844. Bonnet, 1955: 511. 

Araneus seditiosus: — Petrunkevitch, 1911: 314. 
Roewer, 1942: 852. Bonnet (1955: 592) erroneously 
suggests that Petrunkevitch (1911: 314) synony- 
mized Araneus scitulus (Blackwall, 1863) with Ar- 
aneus seditiosus. 

Eustala seditiosa: — Mello-Leitao, 1943: 179. Erro- 
neous transfer. 

Metazygia gressa: — Mello-Leitao, 1943: 187. Erro- 
neous transfer. 

Metepeira gressa: — Levi, 1991: 179. Platnick, 1993: 
449. 

Metepeira seditiosa: — Levi, 1991: 180. Platnick, 1993: 
449. 

Note. Examination of voucher specimens of Vi- 
era suggest that in the behavioral studies of Viera, 
(1986, 1989) and Viera and Costa (1988), tlie name! 
"Metepeira sp. A" is, in fact, M. gressa. 

Description. Female from Punta del Es- 
pinillo, Montevideo, Uruguay. Chelicerae 
brown, lighter on distal inside margins. 
Carapace brown; yellowish white across 
and just behind posterior eye row; median 
white line reaching thoracic furrow, some- 
times thickened into arrow shape (Fig. 
155). Proximal halves of femora, white; re- 
mainder black. Patellae black ventrally, 
white dorsally. Distal halves of tibiae, 
black; remainder white with black annu- 
lation. Femur I with row of four macro- 
setae on anterior side; none on antero- 
ventral side. Dorsal folium white fleur-de- 
lis pattern on black, margined with wavy 
white lines (Fig. 155). Venter black with 
short, wide, white median line; pair of 
small white spots on either side of spiracle 



Metepeira • Piel 55 



(Fig. 156). Sternum black with median 
white hne, often broken (Fig. 156). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.2, anterior laterals 1.3, 
posterior laterals 1.3. Anterior median 
eyes separated by 1.8 diameters, posterior 
median eyes by 1.0, anterior median eyes 
separated from anterior laterals by 2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 3.3 mm. Carapace 1.8 mm long, 
1.4 wide. First femur 1.9 mm, patella and 
tibia 2, metatarsus 1.6, tarsus 0.7. Second 
patella and tibia 1.8 mm, third 1.1, fourth 
1.6. 

Male from Punta del Espinillo, Monte- 
video, Uruguay. Coloration of chelicerae 
and carapace as in female except median 
line brighter, thickened into arrow shape 
(Fig. 153). Leg coloration as in female. Fe- 
mur I with row of four macrosetae on an- 
terior side; five to six on anteroventral side. 
Dorsal folium, venter, and sternum as in 
female (Figs. 153, 154). Ratio of eye di- 
ameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.5, posterior 
laterals 1.4. Anterior median eyes separat- 
ed by 1.3 diameters, posterior median eyes 
by 0.8, anterior median eyes separated 
from anterior laterals by 1.9 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.3 their diameters. Total length 3 mm. 
Carapace 1.8 mm long, 1.3 wide. First fe- 
mur 2.5 mm, patella and tibia 2.3, meta- 
tarsus 1.9, tarsus 0.9. Second patella and 
tibia 1.8 mm, third 1.1, fourth 1.6. 

Diagnosis. The epigynum of M. gressa 
differs from that of M. compso by having 
oval openings (Fig. 151) instead of round 
openings (Figs. 131, 134); it differs from 
M. roraima by being less translucent and 
by having a wider scape (compare Fig. 151 
with Fig. 143). Unlike M. roraima, the em- 
bolus tip in M. gressa is not as bent, and 
the distal apophysis is thinner than the 
point on the embolus where it is attached 
(compare Fig. 149 with Fig. 141). Unlike 
M. compsa, the distal embolic apophysis 
projects away from the embolus tip (Fig. 
149) instead of extending straight from the 



base and parallel to the embolus tip (Figs. 
129, 132). 

Variation. Average body length of 14 fe- 
males examined 4.8 mm, range 3.3 to 6.8 
mm. Average body length of five males ex- 
amined 3.4 mm, range 2.8 to 4.3 mm. 

Natural History. In Uruguay, M. gressa 
live in small colonies of up to five individ- 
uals surrounding the inflorescences of 
Eryngium sp. (Viera and Costa, 1988). The 
number of sticky spirals (14 below the 
hub, 22 above the hub) and radii (c. 40) 
are the same irrespective of age; however 
the length and width of the web differ be- 
tween juveniles (c. 6 and 5 cm, respective- 
ly) and adults (c. 9 and 7 cm, respectively) 
(Viera, 1992). Mature specimens are most 
often collected between September and 
March at low elevation (Fig. 300). 

Distribution. Northern Argentina, Par- 
aguay, and Uruguay (Map 8). 

Records Examined. ARGENTINA Buenos Aires: 
Arrecifes, 34°3'S, 60°7'W, 17.1.1939 (Biraben, MLP); 
Boulogne, 34°30'S, 58°34'W, 15.x. 1938 (Prosen, 
MLP); Colon [?], 33°53'29"S, 61°6'35"W, 20.Lx.l944 
(Torres, MLP); Moreno, 34°39'S, 58°48'W, 15.ii.l966 
(Rossi and Maury, MACN); San Isidro, 34°27'S, 
58°30'W, 15.xii.l937 (Peregra, MACN); San Mi- 
guel—San Fernando, 34°29'S, 58°39'W, 15.vii.l940 
(F. Morrios, MACN); Sierra de la Ventana, 38°9'S, 
61°48'W, 15.iii.l939 (S. H. Bavio, MACN), 15.xi.l954 
(Fritz, MACN), 31.X.1969 (Carlos Grisolia, MCZ), 
15.vii.l972 (Amarrilla, MACN); Tigre, 34°25'S, 
58°34'W (J. M. Viana, MACN). Entre Rios: Concep- 
cion del Uruguay, 32°29'0"S, 58°13'42"W, 4.i.l941 
(Prosen, MLP); Salto Grande, 31°13'S, 57°56'W, 
15.iii.l964 (M. E. Galiano, MEG). Santa Fe: Floren- 
cia Varelo [?], 28°2'S, 59°15'W, 15.xii.l939 (F. Morris, 
MACN). Tucumdn: 30 km S Concepcion, 27°36'S, 
65°35'W, 16.i.l983 (L. E. Pena, AMNH). PARA- 
GUAY Itapiia: San Luis, 27°6'S, 56°36'W, 15.X.1908 
(AMNH). URUGUAY Canelones: Montevideo: De- 
tras del Cerro, Camino de las tropas [?], 34°45'S, 
56°10'W, 6.ii.l963 (R. Capocasale and L. Bmno, 
CAS). Colonia: Punta Gorda^?], 34°28'S, 57°51'W, 
25. ii. 1968 (R. Capocasale and L. Bruno, CAS), 
26.ii.1968 (R. Capocasale and L. Bmno, CAS). Mal- 
donado: Colonia, Cerro de las Animas. Small stones 
in native forest, 34°46'S, 55°19'W, 30.x. 1967 (P San 
Martin, MCZ). Montevideo: Punta del Espinillo, 
34°50'S, 56°26'W, 12.X.1983 (Carmen Viera, MCZ), 
15.xii.l983 (Carmen Viera, CV). Treinta y Tres: Que- 
brada de los Cuervos, 33°10'S, 54°27'W, 27.X.1990 
(Lopez, Perez, Viera, MCZ). 



56 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Metepeira incrassata Group 

Species in the M. incrassata group all 
lack a keel on the median apophysis. Males 
have a distal embolic apophysis that either 
projects toward the embolus tip (e.g.. Fig. 
157), forms a shai-p comer but does not 
actually project forward (e.g.. Fig. 185), or 
forms a smoother, rounded, and gradual 
comer (e.g.. Figs. 192, 206). The M. in- 
crassata species group includes Metepeira 
maya, Metepeira inca, Metepeira gosoga, 
Metepeira ohnec, Metepeira comanche, 
Metepeira pimungan, Metepeira triangu- 
laris, Metepeira arizonica, Metepeira atas- 
cadero, and Metepeira incrassata. 

19. Metepeira maya new species 
Figures 157-163, 317; IViap 14 

Holotype. Male from North Bay, Twin Cays, Stann 
Creek District, Belize, 14.iii.l986, P. Sierwald, in 
USNM. The specific name is a noun in apposition 
after the Indian people of southern Mexico and 
Central America. 

Description. Female paratype from 
Twin Cays, Stann Creek District, Belize. 
Carapace brown with large white eye re- 
gion, white lateral posterior extensions, 
and short median posterior extension (Fig. 
162). Legs white, ringed black on distal 
ends of articles. Femur I with row of four 
macrosetae on anterior side; none on an- 
te roventral side. Dorsum of abdomen with 
usual Metepeira folium, though frequently 
with deep red pigmentation on anterior 
half (Fig. 162). Venter wide, white median 
line with pair of large white spots on either 
side of spiracle (Fig. 163). Sternum with 
wide median white line widening anteri- 
orly with constriction in center (Fig. 163). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 
1.3, posterior laterals 1.3. Anterior median 
eyes separated by 1.2 diameters, posterior 
median eyes by 0.8, anterior median eyes 
separated from anterior laterals by 2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 3.9 mm. Carapace 1.8 mm long, 
1.4 wide. First femur 2.2 mm, patella and 
tibia 2.1, metatarsus 1.6, tarsus 0.8. Sec- 



ond patella and tibia 1.8 mm, third 1.0, 
fourth 1.6. 

Male holotype. Carapace dark with light 
region around eyes and light triangular 
mark anterior to thoracic furrow (Fig. 
160). Legs darker than in female. Femur 
I with row of four macrosetae on anterior 
side; four on anteroventral side. Dorsum, 
venter, sternum as in female except me- 
dian white line on sternum usually broken 
(Figs. 160, 161). Ratio of eye diameters: 
posterior medians and anterior medians 
1.0, anterior laterals 1.4, posterior laterals 
1.2. Anterior median eyes separated by 1.2 
diameters, posterior median eyes by 0.9, 
anterior median eyes separated from an- 
terior laterals by 1.7 diameters of anterior 
lateral eyes, lateral eyes separated by 0.3 
their diameters. Total length 3 mm. Car- 
apace 1.5 mm long, 1.2 wide. First feiTiur 
2.2 mm, patella and tibia 2.3, metatarsus 
1.8, tarsus 0.9. Second patella and tibia 1.8 
mm, third 1.0, fourth 1.4. 

Diagnosis. Unlike others in the M. in- 
crassata species group, M. maya and M. 
inca both have distal embolic apophyses 
that project foiAvard, forming rounded 
bumps (Figs. 157, 164). The male palps of 
these two species differ by the thickness 
of the flagellae and base on the median) 
apophysis: thicker in M. maya than in M. 
inca (compare Fig. 157 with Fig. 164). The 
epigynum of M. maya resembles those of 
M. inca and M. comanche because the oval 
epigynal openings are formed out of mem- 
branous surfaces that are distinctly sepa- 
rate from the base of the scape (Figs. 159, 
166, 187). The epigynal openings of M. 
maya (Fig. 159) are much wider than 
those of M. comanche (Fig. 187); and the 
posterior lobes on M. maya are thickened 
(Fig. 159) but not pointed as in M. inca 
(Fig. 166). 

Variation. Average body length of five 
females examined 5.2 mm, range 3.9 to 6.3 
mm. Average body length of four males 
examined 2.8 mm, range 2.3 to 3.1 mm. 

Natural History. Mature specimens have 
been collected in March through August 



Metepeira • Piel 57 





gressa 
(18) 



It 






maya 
(19) 




A 



r^;''i 




164 



169^^ 170 

inca 
(20) 




Figures 149-156. Metepeira gressa (Keyserling) (sp. 18; 34°50'S, 56°26'W). 149, male palpus, mesal. 150, epigynum, posterior. 

151, epigynum, ventral. 152, male embolic division, ventral. 153, male, dorsal. 154, male, ventral. 155, female, dorsal. 156, 

female, ventral. 

Figures 157-163. Metepeira maya new species (sp. 19; 16°50'N, 88°5'W). 157, male palpus, mesak 158, epigynum, posterior. 

159, epigynum, ventral. 160, male, dorsal. 161, male, ventral. 162, female, dorsal. 163, female, ventral. 

Figures 164-170. Metepeira inca new species (sp. 20; 4°30'S, 81°8'W). 164, male palpus, mesal. 165, epigynum, posterior. 

166, epigynum, ventral. 167, male, dorsal. 168, male, ventral. 169, female, dorsal. 170, female, ventral. 

Scale bars: dorsum and venter figures 1.0 mm. 



58 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



(Fig. 317). Habitats range from mangroves 
at sea level to pine forests at 1,600 m. 

Distribution. Southern Mexico to Costa 
Rica (Map 14). 

Records Examined. BELIZE Stann Creek: Twin 
Cays, Andera Flats, 16°50'N, 88°5'W, 20.iii.l986 (P. 
Sierwald, USNM); Twin Cays, North part of West 
Pond, West of Swamp Doc, 16°50'N, 88°5'W 
28.iii.1986 (P Siei-wald, USNM); Twin Cays, North- 
west Point, North Bay, 16°50'N, 88°5'W, 14.iii.l986 
(P. Sieiw^ald, USNM); Twin Cays, Northwest Point, 
North Bay, ground not flooded, 16°50'N, 88°5'W, 
14.iii.l986 (P Sierwald, USNM); Twin Cays, red 
mangrove, 16°50'N, 88°5'W, 17.V.1985 (Feller, 
USNM); Twin Cays, white mangrove, 16°50'N, 
88°5'W, 15.vi.l984 (Feller, USNM), 5.vi.l985 (Erwin, 
Mathis, Sims, USNM). COSTA RICA San Jose: San 
Jose, 9°56'N, 84°5'W (Tristan & Banks, MCZ). GUA- 
TEMALA Chiquimula: Chiquimula, 14°48'N, 
89°33'W, 22.vii.1947 (C. & P Vaurie, AMNH). Sa- 
catepequez: Antigua, 14°34'N, 90°44'W, 16.viii.l947 
(C. & P Vaurie, AMNH). MEXICO Chiapas: Com- 
itan de Dominguez, 16°1.5'N, 92°8'W, 19.vii.l950 (C. 
J. & M. Goodnight, AMNH); near Rio San Gregorio, 
between Comitan and Ocotal, 15°45'N, 92°0'W, 
18.vii.l950 (C. J. & M. Goodnight, AMNH); pine for- 
est, 15 mi NW Arriaga, 16°25'N, 94°1'W, 27.viii.1966 
(Jean & Wilton Ivie, AMNH); Tuxda Gutierrez, 
16°45'N, 93°7'W, 10.vi.l964 (Palhster, AMNH). NIC- 
ARAGUA Mfltogoipa; Matagaslpa, 12°53'N, 85°57'W 
15.vii.l989 (R. Reinbold, JMM). 

20. Metepeira inca new species 
Figures 164-170, 318; Map 11 

Holotype. Male from Pariiias Valley, Piura, Peru, 
21.V.1939, D. L. & H. E. Frizzell in CAS. The 
specific name is a noun in apposition after the Que- 
chuan people who once luled Peru. 

Description. Female paratype from Par- 
iiias Valley, Piura, Peru. Light reddish 
brown carapace, white around eyes, very 
faint light mark in center (Fig. 169). Legs 
white; slightly darker on dorsal surfaces. 
Femur I with row of four to five macro- 
setae on anterior side; none on anterov- 
entral side. Dorsal folium white; darker on 
shoulders and posteriorly (Fig. 169). Ven- 
ter grayish browii with wide white median 
mark inside U-shaped pattern; pair of 
white spots on either side of spiracle (Fig. 
170). Sternum white (Fig. 170). Ratio of 
eye diameters: posterior medians and an- 
terior medians 0.9, anterior laterals 1.0, 
posterior laterals 1.1. Anterior median 



eyes separated by 1.6 diameters, posterior 
median eyes by 1.3, anterior median eyes 
separated from anterior laterals by 2.5 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 6.8 mm. Carapace 2.9 mm long, 
2.3 wide. First femur 3 mm, patella and 
tibia 3.3, metatarsus 2.6, tarsus 1. Second 
patella and tibia 2.8 mm, third 1.7, fourth 
2.5. 

Male holotype. Light reddish brown 
carapace, white around eyes, faint light tri- 
angular mark in center (Fig. 167). Legs 
white, save for light reddish brown on dor- 
sal surfaces of femora and patellae. Femur 
I with row of three macrosetae on anterior 
side; two to three on anteroventral side. 
Dorsal folium white with wavy black lines 
thickening posteriorly (Fig. 167). Venter 
and sternum as in female (Fig. 168). Ratio 
of eye diameters: posterior inedians and 
anterior medians 1.0, anterior laterals 1.3, 
posterior laterals 1.1. Anterior median 
eyes separated by 1.3 diameters, posterior 
median eyes by 0.8, anterior median eyes 
separated from anterior laterals by 1.9 di- 
ameters of anterior lateral eyes, lateral 
eyes almost touching. Total length 3.7 mm. 
Carapace 1.9 mm long, 1.4 wide. First fe- 
mur 2.9 mm, patella and tibia 2.9, meta-) 
tarsus 2.7, tarsus 1. Second patella and tib- 
ia 2.4 mm, third 1.3, fourth 1.9. 

Diagnosis. Unlike others in the M. in- 
crassata species group, M. inca and M. 
niaija both have distal embolic apophyses 
that project forward, forming rounded 
bumps (Figs. 157, 164). The male palps of 
these two species are separated by the 
thinner flagellae and base on the median 
apophysis of M. inca (compare Fig. 164 
with Fig. 157). The epigynum of M. inca 
resembles those of M. inaija and M. coni- 
anche because the oval epigynal openings 
are formed out of membranous surfaces 
that are distinctly separate from the base 
of the scape (Figs. 159, 166, 187). The epi- 
gynal openings of M. inca (Fig. 166) are 
much wider than those of M. conianche 
(Fig. 187); the posterior lobes on M. inca 



Metepeira • Piel 



59 



are pointed (Fig. 166), not just thickened, 
as in M. may a (Fig. 159). 

Variation. Average body length of six fe- 
males examined 5.7 mm, range 5.2 to 6.8 
mm. Average body length of four males 
examined 3.7 mm, range 2.8 to 4.4 mm. 
Coloration varies significantly among lo- 
calities, especially in the males, where 
some are frequently much darker than 
others. 

Natural History. Mature specimens 
have been collected in April through Oc- 
tober (Fig. 318) at elevations between 300 
and 600 m. 

Distribution. Most northern tip of Peru 
(Map 11). 

Records Examined. PERU Piiira: 12 mi N Man- 
cora, 4°0'S, 80°54'W, ll.xii.l93S; Parinas Valley, 
4°30'S, 81°8'W, 3.iv.l939, 8.iv.l939, 16.iv.l939, 
7.V.1939, 21. V. 1939, 25.vi.1939, 3.vii.l939, 6.viii.l939, 
15.viii.l939, 15.X.1939; Quebrada de Pariiias, 4°32'S, 
81°17'W, 14.iv.l939, 7.V.1939, 21.V.1939; Quebrada 
MogoUon, 4°32'S, 81°4'W, 30.iv.l939, ll.vi.l939, 
18.vi.l939, 21.vi.l939, ll.vii.l939, 16.vii.l939, 
24.ix.1939 (all records: D. L. & H. E. Frizzell, CAS). 

21 . Metepeira gosoga 
Chamberlin and Ivie 

Figures 171-177, 322; Map 9 

Metepeira gosoga Chamberlin and Ivie, 1935: 21 figs 
82-83, 9 . Female holotype from Pilot Knob Valley, 
Mohave Desert, California, in the AMNH, exam- 
ined. Roewer, 1942: 868. Ronnet, 1957: 2820. Levi, 
1977: 200, figs. 28-36. 

Description. Female from Baja Califor- 
nia Norte, Mexico. Carapace diiiy yellow- 
ish brown, lighter anterior half, darker di- 
amond-shaped mark behind eyes (Fig. 
176). Legs same color as carapace; dark 
rings on distal ends of articles. Femur I 
with row of three to five inacrosetae on 
anterior side; two or three macrosetae on 
anteroventral side. Folium lighter than in 
other species, darkening posteriorly (Fig. 
176). Venter black surrounded by yellow. 
Wide white median line with pair of large 
white spots on either side of spiracle. 
Sometimes with parallel pair of lateral 
white lines that join transverse white line 
posteriorly (Fig. 177). Sternum black with 
wide, white line widening anteriorly, often 



broken in center (Fig. 177). Ratio of eye 
diameters: posterior medians and anterior 
medians 0.9, anterior laterals 1.3, posterior 
laterals 1.2. Anterior median eyes separat- 
ed by 1.2 diameters, posterior median eyes 
by 0.6, anterior median eyes separated 
from anterior laterals by 3.4 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.4 their diameters. Total length 8.3 
mm. Carapace 4 mm long, 3.3 wide. First 
femur 4.5 mm, patella and tibia 4.8, meta- 
tarsus 4.5, tarsus 1.3. Second patella and 
tibia 4.3 mm, third 2.5, fourth 3.8. 

Male from Baja California Norte, Mex- 
ico. Male carapace, dorsum, venter, ster- 
num as in female (Figs. 174, 175). Femur 
I with row of five macrosetae on anterior 
side; row of nine macrosetae on anterov- 
entral side. Ratio of eye diameters: poste- 
rior medians and anterior medians 0.8, an- 
terior laterals 1.1, posterior laterals 1.3. 
Anterior median eyes separated by 1 di- 
ameter, posterior median eyes by 0.8, an- 
terior median eyes separated from anterior 
laterals by 2.1 diameters of anterior lateral 
eyes, lateral eyes separated by 0.2 their di- 
ameters. Total length 5.5 mm. Carapace 
2.8 mm long, 2.3 wide. First femur 4.5 
mm, patella and tibia 4.5, metatarsus 4.5, 
tarsus 1.4. Second patella and tibia 3.8 
mm, third 1.9, fourth 3. 

Diagnosis. Within the M. incrassata spe- 
cies group, only M. gosoga, M. inca, and 
M. maya have projecting distal embolic 
apophyses (Figs. 157, 164, 171). Of these, 
only M. gosoga has a distinctly pointed one 
(Fig. 171). The epigynum of M. gosoga dif- 
fers from those of other species in the M. 
incrassata group by the small, oval shape 
of the openings (Fig. 173). 

Variation. Average body length of two 
females examined 6.8 mm, range 5 to 8.5 
mm. Average body length of two males ex- 
amined 4.9 mm, range 4.1 to 5.7 mm. 

Natural History. Mature specimens 
have been collected in June through Au- 
gust (Fig. 322) between 500 and 2,000 m. 
Levi (1977) notes that these have been 
collected on Opuntia and desert vegeta- 
tion. 



60 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Distribution. Southwestern U.S. and 
Baja California Norte, Mexico (Levi, 1977, 
map 1; Map 9). 

Records Examined. MEXICO BaJa Calif. Norte: 23 
mi S Catavina, 29°32'N, 114°57'W, 20.vi.l983 (L. 
Strange & R. Miller, FSCA). USA Arizona: Show 
Low Lake, 34°12'N, 110°0'W, 15.viii.l966 (MCZ). 

22. Metepeira olmec new species 
Figures 178-184, 324; Map 11 

Holotype. Female from Foitin de las Flores, Vera- 
cmz, Mexico, 26.vi.1944, L. L Davis, in AMNH. 
The specific name is a noun in apposition after the 
ancient Indian people of southern Veracruz and Ta- 
basco. 

Description. Female holotype. Carapace 
reddish brown with white region around 
eyes that extends posteriorly along lateral 
margins (Fig. 183). Legs ringed on distal 
ends of articles. Femur I with row of four 
macrosetae on anterior side; none on an- 
teroventral side. Dorsum of abdomen with 
usual Metepeira folium (Fig. 183). Venter 
of abdomen with wide, long white median 
line set within surrounding U-shaped 
markings. Pair of white spots on either 
side of spiracle (Fig. 184). Sternum has 
wide median white line widening anteri- 
orly with broken constriction in center 
(Fig. 184). Ratio of eye diameters: poste- 
rior medians and anterior medians 1.0, an- 
terior laterals 1.1, posterior laterals 1.5. 
Anterior median eyes separated by 1.5 di- 
ameters, posterior median eyes by 0.8, an- 
terior median eyes separated from anterior 
laterals by 2.1 diameters of anterior lateral 
eyes, lateral eyes separated by 0.2 their di- 
ameters. Total length 5.9 mm. Carapace 
2.8 mm long, 2 wide. First femur 2.8 mm, 
patella and tibia 2.8, metatarsus 2.5, tarsus 
1. Second patella and tibia 2.5 mm, third 
1.5, fourth 2.3. 

Male paratype from Fortin de las Flo- 
res, Veracruz, Mexico. Carapace dark with 
light region around eyes and a light mark 
anterior to thoracic furrow (Fig. 181). 
Legs darker than in female. Macrosetae on 
legs variable with spiders size — femur I 
with row of three or four macrosetae on 
anterior side; one to three on anteroven- 



tral side. Dorsum, venter, sternum as in 
female, except median white line on ven- 
ter shortened to spot and median white 
line on sternum often broken (Figs. 181, 
182). Ratio of eye diameters: posterior me- 
dians and anterior medians 1.0, anterior 
laterals 1.3, posterior laterals 1.3. Anterior 
median eyes separated by 1.3 diameters, 
posterior median eyes by 0.8, anterior me- 
dian eyes separated from anterior laterals 
by 1.3 diameters of anterior lateral eyes, 
lateral eyes separated by 0.1 their diame- 
ters. Total length 2.8 mm. Carapace 1.4 
mm long, 1.2 wide. First femur 1.7 mm, 
patella and tibia 1.6, metatarsus 1.1, tarsus 
0.7. Second patella and tibia 1.3 mm, third 
0.8, fourth 1.1. 

Diagnosis. Female M. olmec are diag- 
nosed by their almost perfectly round epi- 
gynal depressions with uniform thickness 
around the posterior edges (Fig. 180). The 
male embolus is relatively short and has a 
vestigial distal embolic apophysis that does 
not project foi^ward, but instead drops off, 
forming a shai^ cuive (Fig. 178). Among 
other species in the M. incrassata species 
group, this embolus shape is also seen in 
M. conianche and M. triangularis: unlike 
M. conianche (Fig. 185), the darker, scler- 
otized portion of the embolus does not ex-\ 
tend over the hump of the distal embolic 
apophysis; unlike M. triangularis (Fig. 
199), the flagellae are thicker than the 
base of the median apophysis. 

Variation. Average body length of four 
females examined 6.2 mm, range 5.6 to 6.7 
mm. Average body length of three males 
examined 2.3 mm, range 2.2 to 2.6 mm. 

Natural History. Mature specimens 
have been collected perennially (Fig. 324) 
at elevations between 500 and 1,400 m. 

Distribution. Montane rain forests from 
Veracruz to Panama (Map 11). 

Records Examined. COSTA RICA San Jose: San 
Antonio de Escanza, 9°59'N, 84°11'W [?], 4.iii.l984 
(W. Eberhard, MCZ). MEXICO Veracruz: Foitin de 
las Flores, 18°54'N, 97°0'W, 26.vi.1944 (L. I. Davis, 
AMNH); Papantla, 20°27'N, 97°19'W, 12.x. 1947 (H. 
Wagner, AMNH). PANAMA Chiriqut: Boquete, 
8°47'N, 82°26'W, 15.viii.l950 (A. M. Chickering, 



Metepeira • Piel 61 





gosoga 
(21) 




184 

olmec 

(22) 




190 ^ 191 

Comanche 
(23) 



Figures 171-177. Metepeira gosoga CnamberWn and Ivie (sp. 21; 29°32'N, 114°57'W). 171, male palpus, mesal. 172, epigynum, 

posterior. 173, epigynum, ventral. 174, male, dorsal. 175, male, ventral. 176, female, dorsal. 177, female, ventral. 

Figures 178-184. Metepeira olmec new species (sp. 22; 18°54'N, 97°0'W). 178, male palpus, mesal. 179, epigynum, posterior. 

180, epigynum, ventral. 181, male, dorsal. 182, male, ventral. 183, female, dorsal. 184, female, ventral. 

Figures 185-191. Metepeira comancfie Levi (sp. 23 [185,188,189] 33°22'N, 99°56'W; [186,187] 25°45'N, 10r55'W; [190, 191] 

26°41'N, 101°23'W). 185, male palpus, mesal. 186, epigynum, posterior. 187, epigynum, ventral. 188, male, dorsal. 189, male, 

ventral. 190, female, dorsal. 191, female, ventral. 

Scale bars: dorsum and venter figures 1 .0 mm. 



62 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



MCZ), 7.viii.l954 (A. M. Chickering, MCZ). Panama: 
Cerro Galera, 8°55'N, 79°38'W, 7.i.l977 (H. W. Levi 
& Y. Lubin, MCZ). 

23. Metepeira comanche Levi 
Figures 185-191, 331; IVlap 11 

Metepeira comanche Levi, 1977: 204, figs. 61-69, S, 
? . Male holotype from 9.7 km west of O'Brien, 
Haskell Co., Texas, in the MCZ, examined. Brig- 
noli, 1983: 275. 

Description. Female from Gloria, Coa- 
huila, Mexico. Carapace dark with light re- 
gion around eyes, extending posteriorly 
behind posterior lateral eyes (Fig. 190). 
Legs ringed. Femur I with row of four ma- 
crosetae on anterior side; one or two on 
anteroventral side. Dorsal folium high in 
contrast; black comma-shaped markings 
shadow white fleur-de-lis pattern (Fig. 
190). Venter of abdomen with wide white 
median line posteriorly set within sur- 
rounding U-shaped marking. Pair of white 
spots on either side of spiracle (Fig. 191). 
Sternum has wide median white line wid- 
ening anteriorly (Fig. 191). Ratio of eye 
diameters: posterior inedians and anterior 
medians 0.9, anterior laterals 1.2, posterior 
laterals 1.1. Anterior median eyes separat- 
ed by 1.2 diameters, posterior median eyes 
by 1.0, anterior median eyes separated 
from anterior laterals by 2.7 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.2 their diameters. Total length 7.8 
mm. Carapace 3.3 mm long, 2.5 wide. 
First femur 3.6 mm, patella and tibia 3.9, 
metatarsus 3.4, tarsus 1.2. Second patella 
and tibia 3.3 mm, third 2, fourth 2.9. 

Male from 9.7 km west of O'Brien, Has- 
kell Co., Texas. Carapace light around 
eyes, darker posteriorly (Fig. 188). Legs 
ringed. Macrosetae on femur I variable; 
usually row of four macrosetae on anterior 
side, six on anteroventral side. Dorsum 
with usual Metepeira folium (Fig. 188). 
Venter with median oval white mark; pair 
of white spots on either side of spiracle 
(Fig. 189). Median white line on sternum 
usually broken (Fig. 189). Ratio of eye di- 
ameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.3, posterior 



laterals 1.3. Anterior median eyes separat- 
ed by 1.4 diameters, posterior median eyes 
by 0.9, anterior median eyes separated 
from anterior laterals by 1.6 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.2 their diameters. Total length 3.1 
mm. Carapace 1.7 mm long, 1.3 wide. 
First femur 2.4 mm, patella and tibia 2.2, 
metatarsus 2, tarsus 0.9. Second patella 
and tibia 1.9 mm, third 1.0, fourth 1.5. 

Diagnosis. The epigynum of M. com- 
anche resembles those of M. may a and M. 
inca because the oval epigynal openings 
are formed out of membranous surfaces 
that are distinctly Separate from the base 
of the scape (Figs. 159, 166, 187). The epi- 
gynum of M. comanche differs from M. 
niaya and M. inca by having much narrow- 
er epigynal openings (compare Fig. 187 
with Figs. 159, 166). The male embolus is 
relatively short and has a vestigial distal 
embolic apophysis that does not project 
forward, but instead drops off, forming a 
sharp curve (Fig. 185). Among other spe- 
cies in the M. incrassata species group, 
this embolus shape is also seen in M. olmec 
and M. triangularis: unlike M. olmec (Fig. 
178), the darker, sclerotized portion of the 
embolus extends over the hump of the dis- 
tal embolic apophysis; unlike M. triangu- 
laris (Fig. 199), the flagellae and base of 
the median apophysis are wide (Fig. 185). 

Natural Histortj. Mature specimens 
have been collected from May to Novem- 
ber (Levi, 1977, Fig. 331). 

Distribution. New Mexico, Texas, and 
northern Mexico (Levi, 1977, map 1; Map 
11). 

Records Examined. MEXICO Coahuila: Gloria, 
26°41'N, 101°23'W, 24.viii.1947 (W. J. Gerisch, 
AMNH); Paila, 25°45'N, 101°55'W, 21.viii.l947 (W. 
J. Gertsch, AMNH). 

24. Metepeira pimungan new species 
Figures 192-198, 330; Map 9 

Holotype. Male from San Miguel Island, Santa Bar- 
bara Co., California, USA, 20.vii.l968, M. E. 
Thompson, in the MCZ. The specific name is a 
noun in apposition after the Indian people who 
once lived on the Channel Islands. 



Metepeira • Piel 



63 



Description. Female paratype from San 
Miguel Island, California USA. Carapace 
with large white eye region and lateral 
posterior extensions. White median arrow- 
shaped mark extends to thoracic furrow 
(Fig. 197). Legs mostly white with rings on 
legs III and IV. Femur I with row of four 
macrosetae on anterior side; one on anter- 
oventral side. Dorsum of abdomen with 
usual Metepeira folium (Fig. 197); venter 
wide, with long white median line, flanked 
by shorter parallel thin white lines on ei- 
ther side. Pair of white spots on either side 
of spiracle. Sternum black with posterior 
drop-shaped white mark (Fig. 198). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.0, anterior laterals 1.4, 
posterior laterals 1.3. Anterior median 
eyes separated by 1.6 diameters, posterior 
median eyes by 1.0, anterior median eyes 
separated from anterior laterals by 3.4 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 6.9 mm. Carapace 3.3 mm long, 
2.6 wide. First femur 3.5 mm, patella and 
tibia 3.5, metatarsus 2.8, tarsus 1.3. Sec- 
ond patella and tibia 3 mm, third 1.9, 
fourth 2.8. 

Male holotype. Carapace, legs, abdo- 
men, venter, sternum as in female, though 
legs darker and not ringed (Figs. 195, 
196). Femur I with row of four to five ma- 
crosetae on anterior side; four to five on 
antero ventral side: both rows concentrated 
distally on femur. Ratio of eye diameters: 
posterior medians and anterior medians 
1.0, anterior laterals 1.3, posterior laterals 
1.3. Anterior median eyes separated by 1.6 
diameters, posterior median eyes by 0.8, 
anterior median eyes separated froin an- 
terior laterals by 1.9 diameters of anterior 
lateral eyes, lateral eyes separated by 0.3 
their diameters. Total length 4.5 mm. Car- 
apace 2.3 mm long, 1.8 wide. First femur 
3.1 mm, patella and tibia 3, metatarsus 2.8, 
tarsus 1. Second patella and tibia 2.5 mm, 
third 1.3, fourth 1.9. 

Diagnosis. Like M. atascadero, but un- 
like other species in the M. incrassata 
group, the posterior end of the sternum of 



M. pimungan has a white water drop mark 
(Figs. 196, 198). Unlike M. atascadero, M. 
pimungan lacks U-shaped white markings 
on the venter, just anterior to the spiracle 
(compare Fig. 198 with Fig. 221). In con- 
trast to other species in the M. incrassata 
group, the embolus on male M. pimungan 
speciinens is curved to forin a gentle S- 
shape (Fig. 192). The female has a squar- 
ish-shaped black mark inside each epigyn- 
al depression (Fig. 198) which is not seen 
in other M. incrassata group species. 

Variation. Average body length of four 
females examined 7.7 mm, range 6.9 to 8.5 
mm. Average body length of five inales ex- 
amined 5.3 mm, range 4.6 to 6.4 mm. 

Natural History. Mature specimens 
have been collected in August (Fig. 330). 
Webs are found near the ground, shel- 
tered from the perennial strong winds by 
Cortjopsis and lupines. 

Distribution. Endemic to San Miguel 
Island (Map 9). 

Records Examined. USA California: San Miguel Is- 
land, 34°2'28.6"N, 120°21'13.6"W, ll.viii.l995"(W. H. 
Piel, MCZ); 34°2'6.1"N, 120°21'9.1"W, 12.viii.l995 
(W. H. Piel, MCZ); 34°2'N, 120°22'W, 20.viii.l968 
(M. E. Thompson, MCZ); 34°3'3.1"N, 120°21'53.3"W, 
13.viii.l995 (W. H. Piel, MCZ); behind dunes on 
beach, 34°2'43.8"N, 120°21'0.8"W, ll.viii.l995 (W. H. 
Piel, MCZ). 

25. Metepeira triangularis (Franganillo) 
Figures 199-205, 337; Map 14 

Mangora triangularis Franganillo, 1930: 21—22. Fe- 
male holotype from Sierra Maestra, lost. Platnick, 
1993: 449. 

Metepeira labijrinthea: — Franganillo, 1936: 75. Bry- 
ant, 1940: 341. Platnick, 1993: 449. 

Metepeira triangidaris: — Archer, 1958: 15, fig. 37, 9 . 

Metepeira acostai Archer, 1958: 15, fig. 36, 9 . Female 
holotype from the savannas, Agramont, Camagiiey 
Province, Cuba, in the AMNH, examined. NEW 
SYNONYMY. 

Metepeira hani Archer, 1965: 132, figs. 11, 17 9, 13 
S . Male and female syntypes from Bani, Domini- 
can Republic, in the AMNH, examined. NEW 
SYNONYMY. Brignofi, 1983: 275. 

Note. Franganillo's collection vials are not la- 
beled, making it impossible to find the holotype. 
Based on the descriptions by Franganillo (1930), 
Archer (1958), the illustrations of Archer (1958, 
1965), and material determined by Archer, I con- 
clude that there is no evidence to support three 



64 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



separate species. For the name M. triangularis, I 
designate a female neotype from Camagiiey Prov- 
ince, Cuba. 21°30'N, 78°10'W, x.l954 (J. T. Acosta, 

AMNH). 

Description. Female from La Descu- 
bierta, Lago Enriquillo, Isla Cabritos, Do- 
minican Republic. Light reddish brown 
carapace, white around eyes (Fig. 204). 
Legs yellowish white, sometimes ringed 
brown at distal ends of articles. Femur I 
with row of three, or sometimes four, ma- 
crosetae on anterior side; none on anter- 
oventral side. Dorsal folium white with 
slightly darker markings posteriorly (Fig. 
204). In some cases, dark anterior marks 
on shoulders of dorsum. Venter of abdo- 
men black with wide white median line, 
sometimes flanked by thinner white lines 
that together form an anchor shape (Fig. 
205). Pair of small white spots on either 
side of spiracle. Sternum black with me- 
dian white line, wider anteriorly (Fig. 205). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 
1.4, posterior laterals 1.4. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 1.1, anterior median eyes 
separated from anterior laterals by 3.3 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 5.9 mm. Carapace 2.7 mm long, 
2 wide. First femur 2.6 mm, patella and 
tibia 2.9, metatarsus 2.5, tarsus 1. Second 
patella and tibia 2.4 mm, third 1.4, fourth 
2.2. 

Male from La Descubierta, Lago Enri- 
quillo, Isla Cabritos, Dominican Republic. 
Carapace as in female, though lighter eye 
region often extends to cover entire ante- 
rior half (Fig. 202). Legs yellowish white. 
Distal half of femora often dark. Femur I 
with row of four macrosetae on anterior 
side; three to five on anteroventral side. 
Dorsal folium white witli slightly darker 
markings posteriorly (Fig. 202). Venter of 
abdomen as in female; sternum black with 
median white line (Fig. 203). Ratio of eye 
diameters: posterior medians and anterior 
medians 1.1, anterior laterals 0.7, posterior 
laterals 1.1. Anterior median eyes separat- 



ed by 1.6 diameters, posterior median eyes 
by 0.8, anterior median eyes separated 
from anterior laterals by 1.0 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.1 their diameters. Total length 4.2 
mm. Carapace 2 mm long, 1.5 wide. First 
femur 2.9 mm, patella and tibia 3, meta- 
tarsus 2.8, tarsus 1.1. Second patella and 
tibia 2.4 mm, third 0.8, fourth 1.3. 

Diagnosis. The epigynum of M. trian- 
gularis differs from other species by the 
thin, sclerotized openings in the shape of 
Ray-Ban sunglasses (Fig. 201). The male 
embolus has a vestigial distal embolic 
apophysis that does not project forward, 
but instead drops off, forming a sharp 
cui-ve (Fig. 199). Among other species in 
the M. incrassata species group, shorter 
versions of this embolus shape is also seen 
in M. Comanche and M. olmec: unlike M. 
Comanche (Fig. 185), the darker, sclero- 
tized portion of the embolus does not ex- 
tend over the hump of the distal embolic 
apophysis; unlike M. olmec (Fig. 178), the 
flagellae and base of the median apophysis 
are much thinner (Fig. 199). 

Variation. Average body length of seven 
females examined 5.5 mm, range 4.6 to 6.3 
mm. Average body length of five males ex- 
amined 3.8 mm, range 3.2 to 4.3 mm. j 

Natural History. Mature specimens 
have been collected from June to Febru- 
ary, although I suspect that like other Ca- 
ribbean species, M. triangularis is actually 
entirely perennial (Fig. 337). 

Distribution. Dominican Republic and 
Cuba (Map 14). 

Records Examined. CUBA Camagiiey Prov.: 
21°30'N, 78°10'W, X.1954 (J. T. Acosta, AMNH). Ma- 
tanzas: Matanzas, Parque Watldns, 23°3'N, 81°35'W, 
9.viii.l955 (A. F. Archer, AMNH); Pan de Palenque 
[?], 23°1'N, 81°43'W, ll.viii.l955 (A. F. Archer, 
AMNH). Oriente: Banes, 20°58'N, 75°43'W, 
2.viii.l955 (A. F Archer, AMNH); Cuabitas, Santiago, 
20°4'N, 75°48'W, 15.xii.l955 (P Alayo, AMNH); 
Puerto Boniato, 20°7'N, 75°47'W, 4.xi.l945 (P Alayo, 
AMNH). Sierra las Casas: Isla de Pinos, 21°53'N, 
82°48'W, 17.viii.l955 (A. F Archer, AMNH). DO- 
MINICAN REPUBLIC Azua: Hatillo, 18°24'N, 
70°32'W, 30.i.l991 (FelLx E. Del Monte, CAS). Ban- 
ahona: Sierra Martin Garcia [?], 18°25'N, 70°30'W, 
8.viii.l958 (A. F Archer & E. de Boyrie Moya, 



Metepeira • Piel 



65 




pimungan 
(24) 




A. 

9 



triangularis 

(25) 




212 

arizonica 
(26) 

Figures 192-198. Metepeira pimungan new species (sp. 24; 34°2'N, 120°22'W). 192, male palpus, mesal. 193, epigynum, 

posterior. 194, epigynum, ventral. 195, male, dorsal. 196, male, ventral. 197, female, dorsal. 198, female, ventral. 

Figures 199-205. Metepeira triangularis (Franganillo) (sp. 25; 18°34'N, 71°44'W). 199, male palpus, mesal. 200, epigynum, 

posterior. 201, epigynum, ventral. 202, male, dorsal. 203, male, ventral. 204, female, dorsal. 205, female, ventral. 

Figures 206-212. Metepeira arizonica Chamberlin and Ivie (sp. 26; 28°53'N, 1 13°4'W). 206, male palpus, mesal. 207, epigynum, 

posterior. 208, epigynum, ventral. 209, male, dorsal. 210, male, ventral. 211, female, dorsal. 212, female, ventral. 

Scale bars: dorsum and venter figures 1 .0 mm. 



66 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



AMNH). La Independencia: 2 km SE El Limon, Ji- 
mani. S side of road, 18°25'N, 71°45'W, 30.i.l991 
(Felix E. Del Monte, MNSD); Isla Cabritos: La Des- 
cubierta, Lago Enriquillo, 18°34'N, 71°44'W, 
18.vi.l981 (E. Marcano, MNSD). Montecristi: Car- 
retera Montecristi, El Morro [?], 19°54'N, 71°39'W, 
2.xii.l991 (EelLx E. Del Monte, MNSD), 21.xii.l991 
(FelLx E. Del Monte, MNSD). Peravia: Las Dunas, 
Prov. Bani [?], 18°25'N, 71°25'W, 23.i.l992 (Felix E. 
Del Monte, MNSD). Frov. Tnijillo Valdez: W of 
Bani, 18°17'N, 70°22'W, 8.viii.l958 (A. F. Archer, 
AMNH). San Juan: 1 km S Las Matas de Farfan, 
18°51'N, 71°31'W, 25.viii.1970 (B. Patterson, MCZ). 

26. Metepeira arizonica 
Chamberlin and Ivie 
Figures 206-213, 319; Map 9 

Metepeira arizonica Chamberlin and Ivie, 1942: 69, 
figs. 182-187, 9,6. Female holotype from Canyon 
Lake, Arizona, in the AMNH, examined. Levi, 
1977: 200, figs. 12, 13, 39^6, 9,3. Brignoh, 1983: 
275. 

Description. Female from Isla Partida, 
Baja Calif. Norte, Mexico. General color- 
ation high in contrast. Carapace dark 
brown; large white eye region, white lat- 
eral posterior extensions, and short median 
posterior extension (Fig. 211). Distinct 
black rings on all legs. Femur I with row 
of four macrosetae on anterior side; four 
on anteroventral side. Dorsum of abdo- 
men with usual Metepeira folium, though 
generally slightly lighter (Fig. 211); venter 
wide, with long white median line set 
within surrounding U-shaped markings. 
Pair of white spots on either side of spi- 
racle (Fig. 212). Wide median white line 
widening anteriorly with constriction in 
center, set on black sternum (Fig. 212). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 
1.4, posterior laterals 1.2. Anterior median 
eyes separated by 1.4 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 2.7 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.1 their diameters. To- 
tal length 6.2 mm. Carapace 2.8 mm long, 
2.2 wide. First femur 3.4 mm, patella and 
tibia 3.5, metatarsus 3, tarsus 1.2. Second 
patella and tibia 3 mm, third 1.9, fourth 
2.8. 



Male from Isla Partida, Baja Calif. Nor- 
te, Mexico. Carapace, legs, abdoiTien, ven- 
ter, sternum as in female, though legs 
darker with less distinct rings, and white 
line on sternum often broken (Figs. 209, 
210). Femur I with row of four macrosetae 
on anterior side; five to eight on anterov- 
entral side. Ratio of eye diameters: poste- 
rior medians and anterior medians 1.1, an- 
terior laterals 1.4, posterior laterals 1.2. 
Anterior median eyes separated by 1.5 di- 
ameters, posterior median eyes by 0.7, an- 
terior median eyes separated from anterior 
laterals by 1.6 diameters of anterior lateral 
eyes, lateral eyes separated by 0.2 their di- 
ameters. Total length 4.4 mm. Carapace 
2.3 mm long, 1.8 wide. First femur 3.3 
mm, patella and tibia 3.3, metatarsus 3.2, 
tarsus 1.2. Second patella and tibia 2.8 
mm, third 1.5, fourth 2.2. 

Diagnosis. Unlike all other species in 
the M. incrassata group, the epigynum of 
M. arizonica is puffy and swollen, causing 
the openings to reduce (from a ventral 
view) to crescent-shaped slits (Fig. 187). 
The embolus differs from others in the M. 
incrassata group because the distal em- 
bolic apophysis is reduced to a gentle arch- 
ing cui-ve (Fig. 206). 

Variation. Average body length of nine 
females examined 5.5 mm, range 4.7 to 6.3 
mm. Average body length of 13 males ex- 
amined 3.9 mm, range 3.1 to 5.1 mm. In 
some males a small ventral keel is visible 
extending beyond the flagella of the me- 
dian apophysis (Levi, 1977, fig. 46). 

'Natural History. Mature specimens are 
most commonly collected between April 
and August (Fig. 319), from dry oak-pine- 
juniper woodland, alfalfa, cactus, and de- 
sert shrub (Levi, 1977). Localities have a 
large range in altitude, from sea level to 
2,300 m; however, spiders from southern 
regions live at higher elevations than spi- 
ders from northern regions (Fig. 213). 

Distribution. American southwest to 
San Luis Potosi (Levi, 1977, map 1; Map 
9). 

Records Examined. MEXICO Baja Calif. Norte: 15 



Metepeira • Piel 



67 











o 


o 






1 


2200 • 


^^ 


° 










B 


1800 - 


° 


^^^ 


o 








o 
1 


1400 ■ 
1000 • 






"""V 


»^ 






i 












■^^ 


° 


1 


600- 
200- 










— ma — a__- 


nn. a. . . . . — ^ 



Latitude (degrees north) 

Figure 213. The elevation of collection localities for M. arizon- 
ica at their corresponding latitudes. Species-specific altitudes 
appear to decrease with distance from the equator. Elevations 
estimated from NOAA database of 5- by 5-minute geographic 
tiles. 



mi S Punta Prieta, 28°58'N, 114°17'W, 7.vii.l973 (S. 
C. Williams & K. B. Blair, CAS); 24 mi S Santa Ines, 
29°20'N, 114°20'W, 7.vii.l973 (S. C. Williams & K. 

B. Blair, CAS); 26 mi S San Felipe, 30°38'N, 
114°50'W, 15.iv.l965 (D. Q. Cavagnaro, C. E. & E. 
S. Ross, V. L. Vesterby, CAS); 6 km NW Racho Santa 
Ines, 29°43'N, 114°43'W, 28.iii.1981 (Paul E. Blom, 
WHO; 6 mi S San Felipe, 30°55'N, 114°52'W 
14.iv.l965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V. 
L. Vesterby, CAS); 9 km NW Racho Santa Ines, 
29°46'N, 114°46'W, 3.vii.l981 (David Crowe, WHC); 
Arroyo de Calamajue at carbonated spring, 29°38'N, 
114°25'W, 26.vi.1973 (S. C. Williams & K. B. Blair, 
CAS); Canon de Cuadalupe, off Laguna Salada, 
32°10'N, 115°48'W, ll.i.l958 (V. Roth, AMNH); El 
Mayor, 32°5'N, 115°13'W, 15.vi.l952 (M. Cazier, W 
Gertsch, & R. Schrammel, AMNH); Isla Mejia, 
29°34'N, 113°35'W, 30.ivl921 (J. C. Chamberlin, 
MCZ); Isla Partida, 28°53'N, 113°4'W, 2.vii.l921 (J. 

C. Chamberlin, MCZ); Isla San Lorenzo, N end, 
28°40'N, 112°52'W, 24.vi.1921 (J. C. Chamberlin, 
CAS); Isla San Lorenzo, south Tobart, 28°36'N, 
112°46'W [?], 9.iv.l921 (J. C. Chamberlin, CAS); 
Puerto Refugio, N end Angel de la Guarda Isl. Webs 
in cracks in cliff overhanging beach, 29°34'N, 
113°32'W, 18.iv.l962 (Howard W. Campbell, CAS); 
San Pedro Martir, 30°45'N, 115°13'W, 18.iv.l921 (J. 
C. Chamberlin, CAS). Coahuila: 25 mi SE San Pedro, 
25°35'N, 102°50'W, 21.viii.l947 (W. J. Gertsch, 
AMNH); Cabos, 25°35'N, 101°43'W, 21.viii.l947 (W. 
J. Gertsch, AMNH); Gloria, 26°41'N, 10r23'W, 
24.viii.1947 (W J. Gertsch, AMNH); Saltillo, 
25°25'N, 101°0'W, 23.V.1952 (M. Cazier, W Gertsch, 
& R. Schrammel, AMNH). Durango: La Loma, 
25°27'N, 103°40'W, 20.viii.l947 (W J. Gertsch, 
AMNH); Providencia, 26°44'N, 105°56'W 
24.viii.1947 (A. M. Davis, AMNH); San Isidro, 60 mi 
NW Durango, 25°1'N, 105°6'W, 19.viii.l947 (W J. 
Gertsch, AMNH). San Luis Potosi: 3 km W Pilares, 
21°55'34"N, 100°48'6"W, 21.X.1994 (W H. Piel, 
MCZ). Sinaloa: Las Saleras Is. (= Isla SaHaca [?]), 
25°11'N, 108°20'W [?], 13.vi.l921 (J. C. Chamberlin, 



CAS). Sonora: 20 mi N Hermosillo, 29°8'N, 
110°58'W, 13.Lx.1966 (Jean & Wilton Ivie, AMNH); 
20 mi SW Sonoyta, 31°38'N, 113°4'W, 13.vi.l952 (W 
J. Gertsch, AMNH); 22 mi E Hermosillo on the 
banks of Rio Sonora, 29°4'N, 110°55'W, 17.viii.l959 
(B. A. Branson, AMNH); Los Angeles, 29°27'N, 
110°46'W, 4.X.1966 (V. Roth, AMNH); Puerto Pefi- 
asco at seashore, 31°20'N, 113°33'W, 3.iv.l968 (D. E. 
Bixler, MCZ). Zacatecas: 14 mi S Fresnillo, 23°5'N, 
102°45'W, 4.viii.l954 (W J. Gertsch, AMNH); 4 mi 
NE Concepcion del Oro, 24°41'N, 101°23'W, 
4.vii.l984 (J. B. Woolley AD). USA Anzona: Santa 
Catalina Mtns., mile 7.9 of Catalina Highway from 
Tuscon to Mt. Lemmon., 32°17'N, 110°48'W, 
19.vi.l985 (W Maddison, MCZ); Tucson, 32°15'N, 
110°57'W, 5.vii.l991 (W H. Piel & G. S. Bodner, 
MCZ). California: Corn Springs, Chuckawalla Mtns., 
10 mi SE Desert Center, 33°33'N, 115°24'W, 
29.xi.1963 (D. C. Lorn, MCZ); Manzanita chaparral, 
San Gabriel Canyon, Coldbrook Ranger Station, 
34°11'N, 117°53'W, 19.viii.l964 (L. Pinter, MCZ). 
Texas: Big Bend Nat'l Park, Old Ranch House on St. 
Elena Rd., 29°10'N, 103°30'W, 25.V.I967 (E. Sabath, 
MCZ). 

27. Metepeira atascadero new species 
Figures 214-221, 336; Map 14 

Holottjpe. Male from San Miguel de Allende, Guana- 
juato, Mexico, 25.x. 1982, George Uetz, in MCZ. 
The specific name is a noun in apposition after the 
Rancho Hotel Atascadero in San Miguel de Allen- 
de. This name was coined by George Uetz when 
he and his colleagues stayed in Rancho Hotel Atas- 
cadero while studying the behavioral ecology of this 
species. This name has been in informal use in the 
literature (e.g., Uetz and Hodge, 1990). 

Description. Female paratype from San 
Miguel de Allende, Guanajuato, Mexico. 
Reddish brown carapace, white around 
eyes, faint light marks extend posteriorly 
behind lateral eyes (Fig. 220). Legs yel- 
lowish, ringed brown at distal ends of ar- 
ticles. Femur I with row of four macro- 
setae on anterior side; three on anterov- 
entral side. Largest branches of fleur-de- 
lis pattern on dorsal folium form large 
paired white spots (Fig. 220). Venter of ab- 
domen black with wide white median line, 
flanked by white U-shape mark (Fig. 221). 
Pair of small white spots on either side of 
spiracle. Sternum black with white poste- 
rior mark (Fig. 221). Ratio of eye diame- 
ters: posterior medians and anterior me- 
dians 1.0, anterior laterals 1.6, posterior 
laterals 1.3. Anterior median eyes separat- 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



ed by 1.4 diameters, posterior median eyes 
by 0.9, anterior median eyes separated 
from anterior laterals by 3.7 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.4 their diameters. Total length 7.5 
mm. Carapace 3.1 mm long, 2.5 wide. 
First femur 4 mm, patella and tibia 4, 
metatarsus 3.9, tarsus 1.3. Second patella 
and tibia 3.3 mm, third 1.8, fourth 2.8. 

Male holotype. Reddish brown cara- 
pace, lighter around eyes, faint light marks 
extend posteriorly behind lateral eyes and 
median eyes (Fig. 218). Legs yellowish 
white, each article gradually turning red- 
dish brown distally. Femur I with row of 
four or five macrosetae on anterior side; 
eight or nine on anteroventral side. Larg- 
est branches of fleur-de-lis pattern on dor- 
sal folium form large paired white spots 
(Fig. 218). Venter of abdomen black with 
wide white median line, flanked by faint 
white U-shape mark (Fig. 219). Pair of 
small white spots on either side of spiracle. 
Sternum black with white posterior mark 
(Fig. 219). Ratio of eye diameters: poste- 
rior medians and anterior medians 1.0, an- 
terior laterals 1.7, posterior laterals 1.4. 
Anterior median eyes separated by 1.8 di- 
ameters, posterior median eyes by 1.0, an- 
terior median eyes separated from anterior 
laterals by 3.6 diameters of anterior lateral 
eyes, lateral eyes separated by 0.5 their di- 
ameters. Total length 5 mm. Carapace 2.5 
mm long, 1.8 wide. First femur 4 mm, pa- 
tella and tibia 4.2, metatarsus 4.5, tarsus 
1.4. Second patella and tibia 3.3 mm, third 
1.7, fourth 2.5. 

Diagnosis. The male embolus of M. 
atascadero has a protruding bump be- 
tween the embolus tip and the basal em- 
bolic apophysis (Figs. 214, 217) which sep- 
arates it from all other Metepeira species. 
As with M. incrasssata (Fig. 224) and, to 
a lesser degree, M. triangularis (Fig. 201), 
the epigynal depressions on either side of 
the scape of M. atascadero (Fig. 216) are 
sclerotized to create a scooped and slick 
quality. The darker epigynal openings in- 
side the depressions are hidden anteriorly 
under the scapes hood. Metepeira atas- 



cadero's epigynum differs from that of M. 
incrassata by its much smaller depressions 
(compare Fig. 216 with 224). 

Variation. Average body length of 12 fe- 
males examined 7.9 mm, range 6 to 9.5 
mm. Average body length of 12 males ex- 
amined 5.7 mm, range 3.5 to 6.9 miTi. 

Natural History. Mature specimens 
have been collected from the end of Au- 
gust to the end of October (Fig. 336). This 
species lives solitarily: only 20% of spiders 
live in aggregations of two or more indi- 
viduals (Uetz and Hodge, 1990). It is 
thought that the low level of social behav- 
ior in M. atascadero occurs because the 
species pursues a risk-prone foraging strat- 
egy (Uetz, 1988a,b). Spiders are found be- 
tween 1,500 and 2,500 m elevation. 

Distribution. Mexican highlands from 
Durango to Guerrero (Map 14). 

Records Examined. MEXICO Coahuila: Saltillo, 14 
mi E in Larrea Desert, 25°25'N, 100°55'W, 
28.vii.1944 (AMNH). Durango: El Taseate, 26°12'N, 
105°7'W, 27.vii.1947 (W. J. Gertsch, AMNH); Yer- 
banis, 80 mi NW Durango, 24°45'N, 103°50'W, 
19.viii.l947 (W. J. Gertsch, AMNH). Guanajuato: 20 
mi E. Guanajuato, 21°1'N, 100°57'W, 15.ix.l976 (C. 

E. Griswold & Jackson, CAS); Guanajuato, 21°1'N, 
101°15'W (N. Banks, MCZ); San Miguel de Allende, 
20°55'N, 100°45'W, 25.x. 1982 (George Uetz, MCZ), 
14.X.1983 (MCZ), 14.X.1985 (MCZ), 17.x.l98^ 
(MCZ), 26.X.1985 (MCZ). Guerrero: Tecalpuico, 25 
km N Iguala, 18°29'N, 99°38'W, l.i.l948 (AMNH). 
Hidalgo: Ozumbilla, 20°9'N, 101°16'W, 2.X.1957 (R. 
Dreisbach, MCZ). Jalisco: Cyarco Onda, 30 km W 
Ojuelos, 21°47'N, 101°53'W (H. Wagner, AMNH). 
Michoacan: Hills N of Patzcuaro, 19°45'N, 101°36'W, 
24.viii.1959 (A. F. Archer, AMNH); Hwy 110, 4 mi 
W Jiquilpan, 19°59'N, 102°47'W, 2.viii.l967 (R. E. 
Leech, REL). Zacatecas: East of Guadalupe, 
22°46'N, 102°31'W, 21.viii.l959 (A. F. Archer, 
AMNH); Guadalupe, 22°45'N, 102°31'W, 16.viii.l947 
(W. J. Gertsch, AMNH). 

28. Metepeira incrassata 

F. O. P. -Cambridge 

Figures 222-228, 323; IVIap 9 

Epeira Salei KeyserUng, 1864: 93, fig., 6,9. Female 
holotype from Oaxaca, Mexico, in BMNH, exam- 
ined. Keyserhng, 1892: 196, fig. 145, 9. Roewer, 
1942: 851. NEW SYNONYMY. 

Metepeira incrassata F. O. R-Cambridge, 1904: 460, 
fig. 11, ?. Female holotype from Jalapa, Mexico, 
in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2821. 



Metepeira • Piel 



69 




220"^" 221 

atascadero 

(27) 



224 





^im^ 227 ' 228 

incrassata 
(28) 




234 



Figures 214-221. Metepeira atascadero new species (sp. 27; 20°55'N, 100°45'W). 214, male palpus, mesal. 215, epigynum, 
posterior. 216, epigynum, ventral. 217, male embolic division, ventral. 218, male, dorsal. 219, male, ventral. 220, female, dorsal. 

221, female, ventral. 

Figures 222-228. Metepeira incrassata F. O. P.-Cambridge (sp. 28 [222,225-228] 19°4'N, 97°2'W; [223,224] 18°54'N, 97°0'W). 

222, male palpus, mesal. 223, epigynum, posterior. 224, epigynum, ventral. 225, male, dorsal. 226, male, ventral. 227, female, 
dorsal. 228, female, ventral. 

Figures 229-235. Metepeira ventura Chamberlin and Ivie (sp. 29; 28°5'N, 114°8'W). 229, male palpus, mesal. 230, epigynum, 
posterior. 231, epigynum, ventral. 232, male, dorsal. 233, male, ventral. 234, female, dorsal. 235, female, ventral. 
Scale bars: dorsum and venter figures 1.0 mm. 



70 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 



Aranea sallei: — F.O.P.-Cambridge, 1904: 519. Roew- 
er, 1942: 851. 

Araneus incrassata: — Petrunkevitch, 1911: 289. 

Araneus sallei: — Petrunkevitch, 1911: 314. 

Metepeira salei: — Levi, 1991: 180. Platnick, 1993: 
449. 

Note. Although M. incrassata is not the oldest 
name, it has been cited more tlian 20 times in the 
general literature (e.g., Caraco et al. 1995; Hieber 
and Uetz, 1990; Hodge and Uetz, 1992, 1995, 
1996; Jakob et al., 1996; Rayor, 1996; Rayor and 
Uetz, 1990, 1993; Uetz, 1988a,b, 1989, 1991, 1992, 
1996; Uetz and Hieber, 1994, 1997; Uetz and Hod- 
ge, 1990; Uetz et al., 1994). In contrast, the senior 
synonym has not been cited outside of taxonomic 
catalogues. In compliance with Article 79 of the 
ICZN (1985), I choose to assert the priority of tlie 
more popular junior name. 

Description. Female from Rancho Chu- 
la-Vista, Veracruz, Mexico. Browii cara- 
pace, darker around margins, lighter 
around eyes and behind lateral eyes (Fig. 
227). Legs brown, yellow at base of articles 
and on articles distal to the femur for legs 
I and II. Femur I with row of three to four 
macrosetae on anterior side; one on anter- 
oventral side. Dorsal folium darker than 
most Metepeira species. White fleur-de-lis 
pattern with thin branches (Fig. 227). Ven- 
ter dark brown to black with white median 
mark that is shorter than it is in most other 
species (Fig. 228). Sternum dark brown to 
black with one or two small white spots, 
usually in the center or anteriorly (Fig. 
228). Ratio of eye diameters: posterior me- 
dians and anterior medians 0.9, anterior 
laterals 1.3, posterior laterals 1.1. Anterior 
median eyes separated by 1.5 diameters, 
posterior median eyes by 1.1, anterior me- 
dian eyes separated from anterior laterals 
by 4.1 diameters of anterior lateral eyes, 
lateral eyes separated by 0.5 their diame- 
ters. Total length 7.8 mm. Carapace 3.6 
mm long, 2.9 wide. First femur 3.9 mm, 
patella and tibia 4.1, metatarsus 3.5, tarsus 
1.4. Second patella and tibia 3.7 mm, third 
2.3, fourth 3.3. 

Male from Rancho Chula-Vista, Vera- 
cruz, Mexico. Carapace, dorsum, venter, 
sternum a darker version of female (Figs. 
225, 226). Base of femora yellow, remain- 
der dark brown; other articles gradually 



turning lighter distally Femur I with row 
of four to six macrosetae on anterior side; 
six to nine on anteroventral side. Ratio of 
eye diameters: posterior medians and an- 
terior medians 1.0, anterior laterals 1.4, 
posterior laterals 1.2. Anterior median 
eyes separated by 1.4 diameters, posterior 
median eyes by 1.0, anterior median eyes 
separated from anterior laterals by 3.7 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.4 their diameters. To- 
tal length 7 mm. Carapace 3.3 mm long, 
2.7 wide. First femur 4.9 mm, patella and 
tibia 5.3, metatarsus 4.8, tarsus 1.7. Sec- 
ond patella and tibia 4.3 mm, third 2.2, 
fourth 3.4. 

Diagnosis. Overall pigmentation of M. 
incrassata is darker than all others in its 
species group. The thin branches on the 
dorsal folium (Figs. 225, 227), the almost 
entirely dark sternum, and the shortened 
median line on the venter (Figs. 226, 228) 
are distinctive. As with M. ohnec (Fig. 
178), M. comanche (Fig. 185), and M. tri- 
angularis (Fig. 199), the embolus of M. in- 
crassata (Fig. 222) curves off sharply from 
a distal embolic apophysis that does not 
project forward. Compared to M. olniec 
(Fig. 178) and M. comanche (Fig. 185), M. 
incrassata's embolus tip beyond the distal 
apophysis is relatively longer and not as 
curved (Fig. 222). Male M. incrassata are 
easily separated from M. triangularis by 
the shape of the median apophysis (com- 
pare Fig. 222 with Fig. 199). As with M. 
atascadero (Fig. 216) and, to a lesser de- 
gree, M. triangularis (Fig. 201), the epi- 
gynal depressions on either side of the 
scape of M. incrassata (Fig. 224) are scler- 
otized to create a scooped and slick qual- 
ity. The darker epigynal openings inside 
the depressions are hidden anteriorly un- 
der the scape's hood. Metepeira incrassa- 
ta's epigynum differs from those of M. 
atascadero and M. triangularis by its much 
larger and disk-shaped depressions (com- 
pare Fig. 224 with Figs. 201, 216). 

Variation. Average body length of 31 fe- 
males examined 7.6 mm, range 6.4 to 9.1 



Metepeira • Piel 



71 



mm. Average body length of 22 males ex- 
amined 6.2 mm, range 4.3 to 8 mm. 

Natural History. Mature specimens 
have been collected in February through 
October but are most frequently found in 
July (Fig. 323). This species lives socially: 
half of all spiders live in aggregations of 
1,000 or more individuals (Uetz and Hod- 
ge, 1990). Colonies are frequently found 
spanning telephone lines, houses, fences, 
and other man-made structures. It is 
thought that this high level of social be- 
havior occurs because the species pursues 
a risk-averse foraging strategy in which the 
variance in survival attenuates with in- 
creasing colony size (Uetz, 1988a,b). Ele- 
vations center around 800 m and range 
from about 500 to 1,500 m. 

Distribution. Tropical Mexico from San 
Luis Potosi to southern Veracruz and Oa- 
xaca (Map 9). 

Records Examined. MEXICO Hidalgo: 20 mi S of 
Jacala, 20°50'N, 99°16'W, 18.iv.l946 (L. I. Davis & 
M. Johnston, AMNH); Chapulhuacan, 21°10'N, 
98°54'W, 20.V.1952 (M. Cazier, W. Gertsch, & R. 
Schrammel, AMNH), 27.vii.1966 (Jean & Wilton 
Ivie, AMNH), 16.vii.l969 (S. & J. Peck, MCZ). Mex- 
ico: 1 mi S Palomas, 19°50'N, 99°5'W, 28.vii.1950 
(AMNH). Oaxaca: Oaxaca, 17°3'N, 96°43'W (Nathan 
Banks, MCZ). Puebla: Huauchinango, 20°11'N, 
98°3'W, 7..X.1947 (H. Wagner, AMNH); north of Xic- 
otepec de Juarez, 20°18'N, 97°57'W 19.ivl967 (W 
B. Peck, MCZ). San Luis Potosi: 10km W Xilitla on 
rl20, 21°22'N, 99°4'W 10.viii.l991 (W H. Piel & G. 
S. Bodner, MCZ). Veracruz: 1 mi SW Tlapacoyan, 
19°57'N, 97°14'W 16.vii.l973 (A. Newton, MCZ); 2 
mi N Fortin de las Flores, 18°56'N, 97°1'W, 
5.viii.l966 (Jean & Wilton Ivie, AMNH); 5 mi E Ori- 
zaba, 18°51'N, 97°4'W 25.vii.1956 (W Gertsch & V. 
Roth, AMNH); Coatepec, 19°27'N, 96°58'W, 
28.vii.1955 (C. & P Vaurie, AMNH), 19.vii.l991 (W. 
H. Piel & G. S. Bodner, MCZ); Fortin de las Flores, 
18°54'N, 97°0'W, 7.vii.l947 (G. & M. Goodnight, 
AMNH), 25.iv.1963 (W. J. Gertsch & W Ivie, 
AMNH), 10.vii.l976 (A. Newton, MCZ), 21.X.1982 
(George Uetz, MCZ), 17.vii.l991 (W. H. Piel & G. S. 
Bodner, MCZ); Jalapa, 19°32'N, 96°55'W, 14.ii.l948 
(H. Wagner, AMNH), 15.iii.l948 (H. Wagner, 
AMNH), 15.X.1962 (N. L. H. Krauss, AMNH); Los 
Naranjos, 18°21'N, 96°10'W 4.iii.l948 (H. Wagner, 
AMNH); near Monte Blanco, 18°58'N, 97°1'W 
3.viii.l973 (A. Newton, MCZ); Orizaba, 18°51'N, 
97°4'W, 6.vii.l963 (D. BLxler, MCZ); Rancho Chula- 
Vista, N of Cordoba, 19°4'N, 97°2'W 18.vii.l991 (W 
H. Piel & G. S. Bodner, MCZ). 



Metepeira ventura Group 

Spiders in the M. ventura group {Me- 
tepeira ventura, Metepeira revillagigedo , 
Metepeira celestun, Metepeira uncata, Me- 
tepeira crassipes, Metepeira chilapae) are 
closely related to those in the M. minima 
group (M. petatlan, M. minima, M. paci- 
fica, M. jamaicensis) . Females in the M. 
ventura group have epigynal openings that 
are wider than long and shaped as trian- 
gles (Fig. 231), ovals (Fig. 252), or squares 
(Figs. 259, 266), with their posterior edges 
open. The posterior edges are observed to 
be open because the posterior lobes at 
their distal edge are wider than the distal 
end of the scape. In many other species, 
the gap between the lobes is narrower 
than the scape and therefore hidden from 
a ventral view, giving the impression that 
the openings form closed shapes (e.g.. Fig. 
123). In addition, with the exception of M. 
celestun (Fig. 245), the scape is relatively 
thin throughout its entire length (Figs. 
231, 238, 252, 259, 266). In males, the em- 
bolus can be slini and elongated with a 
gentle curve (e.g.. Figs. 236, 243), or as 
with some species in the M. minima group, 
it can be tapering to a shaip bend right at 
the tip (e.g.. Figs. 229, 264). In general, 
the larger flagellum is a simple tapering 
extension off the base (Figs. 229, 243, 250, 
257, 264, and to a lesser extent, Fig. 236), 
as opposed to being a distinctly separate 
structure that abruptly cuives off the base 
(e.g.. Fig. 129). Although the flagellae can 
be thin, as in the M. minima species 
group, they are not set off on a distinctly 
narrower stalk (e.g., compare Fig. 264 with 
Fig. 278). 

29. Metepeira ventura 
Chamberlin and Ivie 
Figures 229-235, 320; Map 13 

Metepeira ensenada Chamberlin and Ivie, 1942: 65, 
figs. 166-168, 6. Male holotype from beach near 
Ensenada, Mexico, in the AMNH, examined. Syn- 
onymized by Levi (1977). 

Metepeira ventura Chamberlin and Ivie, 1942: 67, 
figs. 175-179, ?. Female holotype, 1 male and 3 
female paratypes from between Oxnard and Santa 
Monica, California, USA, in the AMNH, examined. 



72 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Levi, 1977: 204, figs. 53-60. Brignoli, 1983: 275. 

Note. As first reviser, Levi (1977) preferred to 
use the name M. ventura because a larger number 
of specimens were available from the type locality. 

Description. Female from 10 mi north 
of Colonia Guerrero, Baja California Nor- 
te, Mexico. Carapace brown with white 
eye region, lateral posterior extensions, 
and white median line (Fig. 234). Legs yel- 
lowish tan with darker rings on fourth pair 
of legs. Femur I with row of four macro- 
setae on anterior side; two on anteroven- 
tral side. Dorsum of abdomen with usual 
Metepeira folium, lighter in anterior third 
(Fig. 234). Venter of abdomen with wide 
white median line posteriorly set within 
hint of surrounding U-shaped marking. 
Pair of white spots on either side of spi- 
racle (Fig. 235). Sternum has wide inedian 
white line widening anteriorly, sometimes 
fragmented (Fig. 235). Ratio of eye di- 
ameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.6, posterior 
laterals 1.4. Anterior median eyes separat- 
ed by 1.4 diameters, posterior median eyes 
by 0.7, anterior median eyes separated 
from anterior laterals by 2.9 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.3 their diameters. Total length 5.5 
mm. Carapace 2.8 mm long, 2.2 wide. 
First femur 3.3 mm, patella and tibia 3.3, 
metatarsus 3, tarsus 1.2. Second patella 
and tibia 2.9 mm, third 1.7, fourth 2.5. 

Male from 10 mi north of Colonia 
Guerrero, Baja California Norte, Mexico. 
Carapace, abdomen, venter as in female 
(Figs. 232, 233). Ringed legs darker than 
in female. Femur I with row of four ma- 
crosetae on anterior side; six to seven on 
anteroventral side. Median white line on 
sternum often broken (Fig. 233). Ratio of 
eye diameters: posterior medians and an- 
terior medians 0.9, anterior laterals 1.2, 
posterior laterals 1.1. Anterior median 
eyes separated by 1.6 diaineters, posterior 
median eyes by 0.8, anterior median eyes 
separated from anterior laterals by 1.9 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 4.3 mm. Carapace 2.2 mm long, 



1.7 wide. First femur 3.6 mm, patella and 
tibia 3.6, metatarsus 3.7, tarsus 1.3. Sec- 
ond patella and tibia 2.8 mm, third 1.5, 
fourth 2.3. 

Diagnosis. Unlike other members of the 
M. minima species group, the larger fla- 
gellum on the median apophysis of M. 
ventura is a tapering extension off the base 
(Fig. 229), as opposed to arising from a 
distinctly separate stalk (Fig. 286). Like M. 
uncata, the epigynal openings of M. ven- 
tura (Fig. 231) are somewhat triangular in 
shape, in contrast to oval (Fig. 288) or slit- 
shaped (Fig. 280) as in other members of 
the M. minima species group. The rela- 
tively shorter scape and the dark circles 
inside the epigynal openings of M. ventura 
(Fig. 231) contrast with M. uncata s rela- 
tively longer scape and the dark sinuous 
shapes inside the epigynal openings (Fig. 
252). 

Variation. Average body length of eight 
females examined 6.7 mm, range 5.6 to 9.7 
mm. Average body length of seven males 
examined 5.1 mm, range 3.9 to 7 mm. 

Natural History. Mature speciinens are 
mostly collected April through September 
(Fig. 320). Altitudes range from near sea 
level to 1,000 m. 

Distribution. California to northwestern 
Mexico (Levi, 1977, map 1; Map 13). 

Records Examined. MEXICO Baja Calif. Not~te: 10 
m E El Rosario, 30°11'N, 115°46'W, 8.vii.l973 (S. C. 
Wilhams & K. B. Blair, CAS); 10 mi N Col. Guerrero, 
28°5'N, 114°8'W, l.b(.1957 (V. Roth, AMNH); 2 mi 
SE Erendira, 3ri9'N, 116°19'W, 12.V.1973 (S. C. 
Wilhams & K. B. Blair, CAS); Isla Cedros, Gran Cafi- 
on, 28°12'N, 115°15'W, 10.iii.l945 (B. F. Osorio Taf- 
all, AMNH); Isla de Cedros, Cerro de Cedros, 
28°12'N, 115°15'W, l.vii.l983 (V. F. Lee, CAS); Isla 
de Cedros, Punta Norte [?], 28°22'N, 115°14'W, 
3.vii.l983 (V. F. Lee, CAS); Isla de Cedros, trail to 
Cerro de Cedros, at spring, 28°12'N, 115°15'W, 
27.Lx.1984 (D. B. Weissman, V. F Lee, CAS); Islas 
San Benito, Benitos del Oeste, 28°18'N, 115°35'W, 
4.vii.l983 (D. C. Lightfoot & V. F Lee, CAS); Islas 
San Benito, Middle Island, 28°19'N, 115°34'W, 
9.iv.l981 (Stanley C. Williams, CAS); Islas San Be- 
nito, South Island, 28°18'N, 115°35'W, 9.iv.l981 
(Stanley C. Williams, CAS); near Consuelo, 6 mi NW 
El Rosario, 30°11'N, 115°46'W 18.iv.l965 (D. Q. 
Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS); 
Rancho Las Parritas, 10 mi S San Quintin, 30°20'N, 



Metepeira • Piel 



73 



115°57'W, 27.vi.1977 (C. E. Griswold, CAS); Santo 
Tomas, 31°33'N, 116°24'W, 8.vii.l953 (W. J. & J. W. 
Gertsch, AMNH). Baja Calif. Sun Bahia de los 
Muertos, 23°58'N, 109°50'W, 20.xii.l958 (H. B. 
Leech, CAS); Desierto del Vizcaino, Laguna Ojo de 
Liebre, sobre frutilla, 27°43'N, 114°15'W, ll.xi.l981 
(A. Cota & M. Jimenez, MLJ); E edge of Sierra Pla- 
ceres, 27°35'N, 114°30'W, 25.iii.1984 (W. J. Pulawsld, 
CAS). Sonora: 6 mi E Navojoa, 27°6'N, 109°23'W, 
23.viii.1965 (W. J. Gertsch & R. Hastings, AMNH). 
USA California: 1 mi NW Winchester (Double 
Butte), 33°43'N, 117°6'W, 7.xii.l976 (W. Icenogle, 
MCZ); Lompoc, by US 63, 34°38'N, 120°27'W, 
17.viii.l966 (L. & P. Pinter, MCZ); Lucia, Monterey 
Co., 36°1'N, 121°33'W, 15.ix.l964 (L. Pinter, MCZ); 
Manzanita chaparral, San Gabriel Canyon, Coldbrook 
Ranger Station, 34°11'N, 117°53'W, 29.V.1965 (L. 
Pinter, MCZ); Santa Catalina Island Area near Hay- 
press Res, 33°23'N, 118°25'W, 6.ii.l993 (Martin C. 
Ramirez & Laura B. Fandino, MCZ); Santa Catalina 
Island, Mt. Torquemada, 33°26'N, 118°33'W, 
15.viii.l965 (L. Pinter, MCZ); Winchester, 33°42'N, 
117°5'W, ll.v.1970 (W. Icenogle, MCZ); Winchester, 
Double Butte, 33°42'N, 117°5'W, 19.V.1974 (W. Icen- 
ogle, MCZ). Nevada: Quin River Crossing, 41°35'N, 
118°27'W, 21.vi.l975 (G. F. Knowlton, MCZ). 

30. Metepeira revillagigedo new species 
Figures 236-242, 301; IVIap 12 

Holotype. Female from south of Isla Socorro, Archi- 
pielago de Revillagigedo, State of Colima, Me.xico, 
15.xii.l988, M. Jimenez, in MCZ. The specific 
name is a noun in apposition after the locality. 

Description. Female holotype. Carapace 
dark brown with light region in median 
eye quadrangle and surrounding lateral 
eyes. Pair of darker, elliptic, walnut leaf 
shapes on lighter patch near center of car- 
apace (see Fig. 241). Darker rings on distal 
ends of femora, patellae, tibiae, and prox- 
imal dorsal portion of tibia I, II. Femur I 
with three to four macrosetae on anterior 
side; none on the anteroventral side. Dor- 
sum of abdomen with white oak leaf foli- 
um, margined with dark markings, partic- 
ularly on trailing edges of lobes (Fig. 241). 
Coloration has a slightly golden hue. Dark 
lateral band wraps around sides of abdo- 
men and stretches up over anterior dorsal 
portion. Venter with two white spots on 
either side of spiracle. Anterior to spiracle, 
slight indication of a V-shaped mark with 
longitudinal extensions reaching halfway 
up abdomen. Wide, white medial longitu- 



dinal line to epigynal groove (Fig. 242). 
Sternum with wide, white longitudinal 
mark widening anteriorly (Fig. 242). Ratio 
of eye diameters: posterior medians and 
anterior medians 0.8, anterior laterals 1.1, 
posterior laterals 1. Anterior median eyes 
separated by 1.3 diameters, posterior me- 
dian eyes by 1.0, anterior median eyes sep- 
arated from anterior laterals by 2.5 diam- 
eters of anterior lateral eyes, lateral eyes 
separated by 0.3 their diameters. Total 
length 7.5 mm. Carapace 3.2 mm long, 2.5 
wide. First femur 3.5 mm, patella and tibia 
3.8, metatarsus 3.2, tarsus 1.2. Second pa- 
tella and tibia 3.2 mm, third 1.8, fourth 
2.8. 

Male paratype froin Isla Socorro, Coli- 
ma. Carapace dirty yellowish brown with 
lighter posteriorly pointing acute triangle 
in center (Fig. 239). Legs same color as 
carapace, except lighter on proximal half 
of femora. Femur I with row of four ma- 
crosetae on anterior side, row of two ma- 
crosetae on anteroventral side. Male ab- 
domen lighter and with less contrast than 
female (Fig. 239). Venter with a much re- 
duced and shorter longitudinal line than 
female. Sternum as in female, except lon- 
gitudinal line more often broken (Fig. 
240). Ratio of eye diameters: posterior me- 
dians and anterior medians 1.0, anterior 
laterals 1.1, posterior laterals 1.1. Anterior 
median eyes separated by 1.5 diameters, 
posterior median eyes by 0.8, anterior me- 
dian eyes separated from anterior laterals 
by 1.4 diameters of anterior lateral eyes, 
lateral eyes separated by 0.3 their diame- 
ters. Total length 3.2 mm. Carapace 1.7 
mm long, 1.1 wide. First femur 2.5 mm, 
patella and tibia 2.5, metatarsus 2.3, tarsus 
0.9. Second patella and tibia 2 mm, third 
1.0, fourth 1.5. 

Diagnosis. The epigynum of M. revilla- 
gigedo looks veiy different from those of 
other species in the M. ventura group. In- 
stead of having small, sharply delineated 
depressions on either side of the scape 
(e.g.. Fig. 259), M. revillagigedo has larger 
but more gradual depressions (Fig. 238). 
Despite the unique appearance of the epi- 



74 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



gynum, the affinity between M. revillagi- 
gedo and the M. ventura species group can 
be seen in the black comma-shaped marks 
inside the epigynal depressions (Fig. 238) 
similar to those in, for example, M. Ven- 
tura (Fig. 231). The male emboli of M. 
revillagigedo (Fig. 236) and M. celestun 
(Fig. 243) are veiy similar in shape and 
relatively longer than in other species in 
the M. Ventura group. Males of M. revil- 
lagigedo differ from males of M. celestun 
by the thickness and shape of the median 
apophysis (compare Fig. 236 with Fig. 
243). 

Natural History. Specimens have only 
twice been collected: once in December 
and once in April (Fig. 301). This species 
is known to live in fig trees. 

Distribution. This species is found on 
Isla Socorro in the Pacific Ocean (Map 
12), the island being one of several that 
form the Archipielago de Revillagigedo. It 
is well isolated, situated about 465 km 
south of Baja California Sur and 588 km 
west of the Jalisco coastline. 

Records Examined. MEXICO Colinia: Archipiela- 
go de Revillagigedo: Isla Socorro, 18°45'N, 110°57'W, 
26.iv.1932 (Templeton, Crocker Exped., CAS); Ar- 
chipielago de Revillagigedo: Sur de la Isla Socorro, 
18°44'N, 110°57'W, 15.xii.l988 (M. Jimenez, MCZ). 

31 . Metepeira celestun new species 
Figures 243-249, 326; IVlap 13 

Holotype. Male from Celestun, Yucatan, Mexico, 
24.vii.1991, W. H. Piel & G. S. Bodner, in MCZ. 
The specific name is a noun in apposition after the 
locality. 

Note. Since males of this species were never 
found witli females, one cannot be sure absolutely 
that they are conspecific. Several facts argue for 
conspecificity: all specimens were collected in sim- 
ilar habitats and during the same season; dorsal fo- 
lium of males and females share similar patterns 
and both have hint of gold coloration; based on 
parallel correspondence widi other species, die 
male palp is arguably compatible with the female s 
epigynum. 

Description. Female paratype from 
Edzna, Campeche, Mexico. Carapace yel- 
lowish tan with lighter area around eyes 
(Fig. 248). Legs yellowish tan with darker 
rings on fourth pair of legs. Femur I with 



row of four macrosetae on anterior side; 
sometimes four on anteroventral side. 
Dorsal folium as in other Metepeira, ex- 
cept speckled an unusual reddish gold col- 
or (Fig. 248). Venter wide white median 
line with pair of large white spots on either 
side of spiracle (Fig. 249). Sternum has 
wide median white line widening anteri- 
orly with constriction in center (Fig. 249). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 

1.3, posterior laterals 1.2. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated froin anterior laterals by 2.4 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 5.2 mm. Carapace 2.5 mm long, 
1.9 wide. First femur 2.6 mm, patella and 
tibia 2.8, metatarsus 2.2, tarsus 0.9. Sec- 
ond patella and tibia 2.4 mm, third 1.4, 
fourth 2. 

Male holotype. Carapace dark with light 
region around eyes and a light triangular 
mark anterior to thoracic furrow (Fig. 
246). Coxae and proximal third of femora 
white, remaining two-thirds black; other 
articles similarly ringed. Femur I with row 
of four macrosetae on anterior side; three 
on anteroventral side. Dorsal folium with 
slight gold iridescence. Fleur-de-lis pat- 
tern leafy and pinnate (Fig. 246). Venter, 
sternum as in female, except median white 
line on sternum often broken (Fig. 247). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.1, anterior laterals 

1.4, posterior laterals 1.3. Anterior median 
eyes separated by 1.6 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 1.4 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 2.8 mm. Carapace 1.4 mm long, 
1.1 wide. First femur 1.8 mm, patella and 
tibia 1.8, metatarsus 1.4, tarsus 0.8. Sec- 
ond patella and tibia 1.5 mm, third 0.8, 
fourth 1.2. 

Diagnosis. The overall coloration of M. 
celestun in alcohol has an unusually golden 
quality that is rare among preserved Me- 



Metepeira • Piel 75 




Figures 236-242. Metepeira revillagigedo new species (sp. 30; 18°44'N, 110°57'W). 236, male palpus, mesal. 237, epigynum, 

posterior. 238, epigynum, ventral. 239, male, dorsal. 240, male, ventral. 241, female, dorsal. 242, female, ventral. 

Figures 243-249. Metepeira celestun new species (sp. 31 [243,246,247] 20°56'N, 90°21'W; [244,245,248,249] 19°35'N, 

90°15'W). 243, male palpus, mesal. 244, epigynum, posterior. 245, epigynum, ventral. 246, male, dorsal. 247, male, ventral. 

248, female, dorsal. 249, female, ventral. 

Figures 250-256. Metepeira uncata F. O. P.-Cambridge (sp. 32 [250,251,253-256] 14°40'N, 92°9'W; [252] 14°49'N, 91°31'W), 

250, male palpus, mesal. 251, epigynum, posterior. 252, epigynum, ventral. 253, male, dorsal. 254, male, ventral. 255, female, 

dorsal. 256, female, ventral. 

Scale bars: dorsum and venter figures 1.0 mm. 



76 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



tepeira. Unlike all other members of the 
M. Ventura species group, the scape of M. 
celestun is very wide at its base, thereby 
partly concealing the epigynal openings 
(Fig. 245). However, if the scape is dis- 
carded, the exposed openings have black 
comma-shaped marks similar to those of 
M. Ventura (Fig. 231), thereby confirming 
the species' affinities with other members 
in the M. ventura species group. Unlike 
other members, the posterior lobes nearly 
touch each other near their base (Fig. 
244). The emboli of M. celestun and M. 
revillagigedo are very similar in shape and 
relatively longer than in other species in 
the M. ventura group (Fig. 243). Metepei- 
ra celestun differs from M. revillagigedo by 
the thickness and shape of the median 
apophysis (compare Fig. 243 with Fig. 
236). 

Variation. Average body length of four 
feinales examined 5.6 mm, range 5.2 to 6.4 
mm. Average body length of three males 
examined 2.7 mm, range 2.2 to 2.9 mm. 
Female scape sometimes wider than it ap- 
pears in Fig. 245, resembling Metepeira 
arizonica. Legs and carapace color vary 
from yellowish tan to orange-red. 

Natural History. Mature specimens 
have been collected in July (Fig. 326) in 
forested clearings, roadside bushes, palms, 
and swampy areas near the beach. 

Distribution. Yucatan peninsula (Map 
13). 

Records Examined. MEXICO Campeche: Edzna, 
19°35'N, 90°15'W, 22.vii.1991 (W. H. Piel & G. S. 
Bodner, MCZ). Quintana Roo: Kohunlich ruins, 9 km 
S Franciso Villa, 18°26'N, 88°48'W, 15.vii.l983 (R. S. 
Anderson, MCZ). Yucatan: 3km S San Felipe, 
21°32'N, 88°14'W, 25.vii.1991 (W. H. Piel & G. S. 
Bodner, MCZ); 4 km N Xocenpich, 12 km N Piste, 
on road to Dzitas, 20°47'N, 88°34'W, 20.vii.l983 (W. 
Maddison, MCZ); Balankanche Cave, 2 km E Chi- 
chen Itza, 20°40'N, 88°33'W, 19.vii.l983 (W. Mad- 
dison, MCZ); beach north of Celestun, 20°56'N, 
90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner, 
MCZ); Chichen Itza ruins on HWY 180, seasonal for- 
est, 20°40'N, 88°34'W, 19.vii.l983 (W. Maddison & 
R. S. Anderson, MCZ); Uxmal, 20°22'N, 89°46'W, 
23.vii.1991 (W. H. Piel & G. S. Bodner, MCZ). 



32. Metepeira uncata 
F. O. P. -Cambridge 
Figures 250-256, 329; Map 12 

Metepeira uncata F. O. P.-Cambridge, 1903: 459, fig. 
8, 6. Male holotype from Santa Ana, Guatemala, 
in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2823. 

Araneus uncatus: — Petrunkevitch, 1911: 321. 

Description. Female from Ayutla, San 
Marcos, Guatemala. Carapace dark brown 
with light region surrounding the eyes and 
extending posteriorly behind the lateral 
eyes. Pair of darker feather-shaped patches 
anterior of thoracic furrow, touching pos- 
teriorly to create a U-shape (see Fig. 255). 
Dark rings on distal half of leg articles, an- 
terior side of patellae, and dark markings 
on dorsal, proximal end of tibia I and II. 
Femur I with four macrosetae on anterior 
side; one on anteroventral side. Dorsum of 
abdomen with white fleur-de-lis folium; 
this pattern thinner than in most other 
species. Fleur-de-lis pattern on a dark 
background, outlined on either side by 
thin white stripe (Fig. 255). Dark lateral 
band follows sides of abdomen and 
stretches up over anterior dorsal portion. 
Venter with two white spots on each side 
of spiracle; wide, white, medial longitudi- 
nal line starts anterior to the colulus and 
ends posterior to the epigynal groove. 
Thin, faint lines run parallel to wide me- 
dian one anterior to white spots. White 
patch separates epigynum and dark de- 
pression posterior to pedicel (Fig. 256). 
Sternum black with white dewdrop- 
shaped mark at posterior end (Fig. 256). 
Ratio of eye diameters: posterior medians 
and anterior medians 1.0, anterior laterals 
1.3, posterior laterals 1.2. Anterior median 
eyes separated by 1.6 diameters, posterior 
median eyes by 1.1, anterior median eyes 
separated froin anterior laterals by 4 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.4 their diameters. To- 
tal length 7.8 mm. Carapace 3.7 mm long, 
2.9 wide. First femur 4 mm, patella and 
tibia 4.3, metatarsus 3.8, tarsus 1.4. Sec- 
ond patella and tibia 3.6 mm, third 2.3, 
fourth 3.2. 



Metepeira • Piel 77 



Male from Ayutla, San Marcos, Guate- 
mala. Carapace dirty yellowish brown with 
lighter mark in center (Fig. 253). Saine 
color as carapace except lighter on proxi- 
mal half of femora. Femur I with row of 
nine macrosetae on anterior side; five on 
anteroventral side. Male abdomen similar 
to female, but leaves of fleur-de-lis pattern 
thinner (Fig. 253). Venter with a shorter 
longitudinal line, as compared with female 
(Fig. 254). Sternum with posterior white 
dewdrop as in female, and additional an- 
terior white mark near labium (Fig. 254). 
Ratio of eye diameters: posterior medians 
and anterior medians 0.9, anterior laterals 
1.3, posterior laterals 1.3. Anterior median 
eyes separated by 1.5 diameters, posterior 
median eyes by 1.0, anterior median eyes 
separated from anterior laterals by 2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 4.5 mm. Carapace 2.3 mm long, 
1.8 wide. First femur 3.4 mm, patella and 
tibia 3.4, metatarsus 3.2, tarsus 1.1. Sec- 
ond patella and tibia 2.7 mm, third 1.4, 
fourth 2. 

Diagnosis. The generally darker pig- 
mentation (Fig. 255) and the small white 
mark on the sternum (Fig. 256) separate 
M. uncata from all other members of the 
M. Ventura species group. Like M. ventura 
(Fig. 231) but unlike other members of 
the M. ventura species group, the open- 
ings to the epigynum are more triangular 
to oval (Fig. 252), rather than square (Figs. 
259, 266). Also distinctive are the sinuous 
black lines that form upside-down U- 
shapes inside each epigynal opening (Fig. 
252), in contrast to the comma-shaped 
marks in M. ventura or M. chilapae (Figs. 
231, 266). Like M. chilapae but unlike oth- 
er members of the M. ventura group, the 
median apophysis and flagellae of M. un- 
cata are slimmer and of uniform thickness 
(Figs. 250, 264). The palp of M. uncata 
differs from that of M. chilapae by the 
more basal position of the bend in the em- 
bolus tip (compare Fig. 250 with Fig. 264). 

Variation. Average body length of five 
females examined 8.2 mm, range 7.5 to 8.8 



mm. Average body length of four males 
examined 5.6 mm, range 4.5 to 6.4 mm. 

Natural Historij. Mature specimens 
have been collected in August (Fig. 329). 

Distribution. Southwestern Guatemala 
to northern Costa Rica (Map 12) at alti- 
tudes ranging from 100 to 3,000 m. 

Records Examined. COSTA RICA Cordillera: 20 
km N Siquires, 10°9'N, 84°17'W, 15.viii.l980 (W. 
Eberhard, MCZ). GUATEMALA El Quiche: Chichi- 
castenango, 14°56'N, 91°7'W, 6.viii.l947 (C. & P. 
Vaurie, AMNH). Huehuetenango: Todos Santos Cu- 
chumatan, 15°31'N, 91°37'W, 16.viii.l979 (C. E. 
Griswold, CAS). Quetzaltenango: El Baul, 1 km S 
Quetzaltenango, 14°49'N, 91°31'W, 14.viii.l979 (T C. 
Meikle & C. E. Griswold, CAS); Quetzaltenango, 
14°50'N, 91°31'W, 16.viii.l950 (C. J. & M. Good- 
night, AMNH). San Marcos: Ayutla, 14°40'N, 92°9'W, 
19.viii.l947 (AMNH). 

33. Metepeira crassipes 
Chamberlin and Ivie 
Figures 257-263, 309; Map 12 

Metepeira josepha Chamberlin and Ivie, 1942; 64, fig. 
165, 9 . Female holotype from Kings Mtn. near 
Palo Alto, California in the AMNH. Synonymized 
by Levi (1977). 

Metepeira crassipes Chamberlin and Ivie, 1942: 66, 
figs. 171-173, 9, S. Male holotype, female, male 
paratypes from Laguna Beach, California in the 
AMNH. Levi, 1977: 202, figs. 47-52, 9,6. 

Note. As first reviser, Levi (1977) preferred to 
use name M. crassipes. 

Description. Female from Isla San Pe- 
dro Nolasco, Sonora, Mexico. Carapace 
with large white eye region and lateral 
posterior extensions (Fig. 262). Legs 
ringed at distal ends of articles — though 
sometimes only lightly. Femur I with four 
macrosetae on anterior side aligned in 
straight row; one to four light setae on an- 
teroventral side. Dorsum of abdomen with 
usual Metepeira folium (Fig. 262); venter 
wide, with long white median line set 
within U-shaped thinner white lines. Pair 
of white spots on either side of spiracle 
(Fig. 263). Sternum black with wide me- 
dian white line widening anteriorly (Fig. 
263). Ratio of eye diameters: posterior me- 
dians and anterior medians 1.1, anterior 
laterals 1.4, posterior laterals 1.3. Anterior 
median eyes separated by 1.4 diameters. 



78 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



posterior median eyes by 0.8, anterior me- 
dian eyes separated from anterior laterals 
by 3 diameters of anterior lateral eyes, lat- 
eral eyes separated by 0.1 their diameters. 
Total length 5.7 mm. Carapace 2.8 mm 
long, 2.1 wide. First femur 3.2 mm, patella 
and tibia 3.1, metatarsus 2.8, tarsus 1. Sec- 
ond patella and tibia 2.7 mm, third 1.6, 
fourth 2.5. 

Male from Winchester, California. Car- 
apace as in female except often with me- 
dian white mark. Femur I with row of four 
macrosetae on anterior side; about four to 
eight on anteroventral side. Dorsum, ven- 
ter as in female (Fig. 260); median white 
line on sternum more often broken (Fig. 
261). Ratio of eye diameters: posterior me- 
dians and anterior medians 1.0, anterior 
laterals 1.6, posterior laterals 1.4. Anterior 
median eyes separated by 1.2 diameters, 
posterior median eyes by 0.6, anterior me- 
dian eyes separated from anterior laterals 
by 2 diameters of anterior lateral eyes, lat- 
eral eyes separated by 0.3 their diameters. 
Total length 4 mm. Carapace 2.1 mm long, 
1.5 wide. First femur 3.2 mm, patella and 
tibia 3.2, metatarsus 3.2, tarsus 1.1. Sec- 
ond patella and tibia 2.5 mm, third 1.3, 
fourth 2. 

Diagnosis. Unlike other species in the 
M. Ventura species group, the openings to 
the epigynum of M. crassipes resemble 
those of M. chilapae because they are 
shaped like squares with rounded edges 
(compare Fig. 259 with Fig. 266). How- 
ever, compared to M. chilapae, the epigyn- 
al openings of M. crassipes are relatively 
smaller and the scape is relatively longer. 
The embolus of M. crassipes is distin- 
guished by its gentle cui'vature (Fig. 257), 
in contrast to much more abi"upt cui"vature 
seen in other species in the M. ventura 
group (e.g.. Fig. 264), with the exception 
of M. revillagigeclo. The median apophysis 
of M. crassipes (Fig. 257) is slimmer than 
that of M. revillagigedo (Fig. 236). 

Variation. Average body length of four 
females examined 6 mm, range 5 to 6.5 
mm. Average body length of two males ex- 
amined 4.4 mm, range 4.1 to 4.7 mm. 



Sometimes the scape is greatly swollen, 
adding to its relative width. In such cases 
the scape can cover the epigynal openings, 
which may mislead the investigator to con- 
fuse it for Metepeira arizonica. 

Natural Histortj. Levi (1977) reports 
that males in the U.S. have been collected 
from April to October, primarily in Cali- 
fornia buckwheat and sage. Mexican re- 
cords expand this seasonality to include 
the entire year (Fig. 309). 

Distribution. From northern California 
in the U.S. to Baja California Sur, Mexico 
(Levi, 1977, map 1; Map 12). 

Records Examined. MEXICO Baja Calif. Norte: 
Santo Tomas, 31°33'N, 116°24'W, 8.vii.l953 (W. J. & 
J. W. Gertsch, AMNH), 12.xi.l976 (S. C. Williams & 
K. B. Blair, CAS). Baja Calf. Sun 26 mi S Loreto, 
25°37'N, 111°17'W, l.i.l977 (C. E. Griswold & L. 
Vincent, CAS); Isla Magdalena, Puerto Magdalena, 
24°38'N, 112°9'W, 16.iii.l957 (R. Zweifel, AMNH); 
Isla San Francisco, South Side, 24°50'N, 110°35'W, 
19.V.1970 (S. C. Williams & V. F. Lee, CAS); Sierra 
San Nicolas, 26°32'N, 111°36'W [?] (Eisen & Vaslit, 
MCZ). Sonora: Isla San Pedro Nolasco, 27°58'N, 
111°25'W, 17.iv.l921 (G. C. Chamberlin, MCZ); Si- 
erra de Alamos, 30°51'N, 112°2'W [?], 19.i.l968 (V. 
Roth, AMNH). USA California: 1 mi NW Winches- 
ter (Double Butte), in web between Artemisia cali- 
fornica bushes, 33°43'N, 117°6'W, 4.xii.l976 (W. 
Icenogle, MCZ). 

34. Metepeira chilapae 
Chamberlin and Ivie 
Figures 264-270, 333; Map 13 

Metepeira chilapae Chamberlin and Ivie, 1936: 45, 
figs. 119-121, S. Male holotype from Chilapa, 
Guerrero, Mexico, in the ZMB. Roewer, 1942: 868. 

Metepeira chilapica: — Bonnet, 1957: 2820. Unjusti- 
fied eiuendation. 

Note. Examination of Chamberlin and Ivies 
(1936) and Levi's (personal illustrations) figures of 
the holotype was sufficient to identify M. chilapae 
accurately. 

Description. Female from Cocoyoc, 
Morelos, Mexico. Brownish black cara- 
pace, paler around and just posterior to 
lateral eyes (Fig. 269). Distal halves of 
ventral leg articles black, elsewhere yel- 
lowish. Femur I with row of four macro- 
setae on anterior side; zero to three setae 
on anteroventral side. Dorsal folium a 
white fleur-de-lis marking set on golden- 



Metepeira • Piel 79 




257 



crassipes 
(33) 



260 261 




269 

chilapae 
(34) 




270 



276 




277 



272 



petatlan 
(35) 



Figures 257-263. Metepeira crassipes Chamberlin and Ivie (sp. 33 [257-261] 33°43'N, 117°6'W; [262,263] 27°58'N, 1 1 1°25'W). 

257, male palpus, mesal. 258, epigynum, posterior. 259, epigynum, ventral. 260, male, dorsal. 261, male, ventral. 262, female, 

dorsal. 263, female, ventral. 

Figures 264-270. Metepeira cliilapae Chamberlin and Ivie (sp. 34 [264] 17°39'N, 99°22'W; [265-270] 17°39'N, 99°22'W). 264, 

male palpus, mesal. 265, epigynum, posterior. 266, epigynum, ventral. 267, male, dorsal. 268, male, ventral. 269, female, dorsal. 

270, female, ventral. 

Figures 271-277. Metepeira petatlan new species (sp. 35 [271,274,275] 17°14'N, 100°53'W; [272,273,276,277] 17°17'32"N, 

101°2'40"W). 271, male palpus, mesal. 272, epigynum, posterior. 273, epigynum, ventral. 274, male, dorsal. 275, male, ventral. 

276, female, dorsal. 277, female, ventral. 

Scale bars: dorsum and venter figures 1.0 mm. 



80 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



brown speckled pattern over white. Foli- 
um darkens posteriorly (Fig. 269). Ante- 
rior shoulders black. Venter black with 
wide median white line and pair of white 
spots on eidier side of spiracle (Fig. 270). 
Sternum brownish black with wide, white 
line widening anteriorly (Fig. 270). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.0, anterior laterals 1.1, 
posterior laterals 1.1. Anterior median 
eyes separated by 1.3 diameters, posterior 
median eyes by 0.8, anterior median eyes 
separated from anterior laterals by 2.2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.2 their diameters. To- 
tal length 5.8 mm. Carapace 2.7 mm long, 
2.3 wide. First femur 2.9 mm, patella and 
tibia 3, metatarsus 2.6, tarsus 1. Second 
patella and tibia 2.4 mm, third 1.6, fourth 
2.3. 

Male from 6 mi northeast of Tixtla de 
Guerrero, Guerrero, Mexico. Carapace, 
dorsum, venter, sternum darker and more 
contrasty version of female (Figs. 267, 
268). Distal halves of ventral leg articles 
black, elsewhere yellowish. Femur I with 
row of four macrosetae on anterior side; 
three on an tero ventral side. Ratio of eye 
diameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.3, posterior 
laterals 1.3. Anterior median eyes separat- 
ed by 1.7 diameters, posterior median eyes 
by 0.9, anterior median eyes separated 
from anterior laterals by 1.9 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.3 their diameters. Total length 3.5 
mm. Carapace 1.8 mm long, 1.4 wide. 
First femur 2.3 mm, patella and tibia 2.1, 
metatarsus 1.8, tarsus 0.8. Second patella 
and tibia 1.8 mm, third 0.9, fourth 1.4. 

Diagnosis. The openings to the epigyn- 
um resemble those of M. crassipes be- 
cause they are shaped like squares with 
rounded edges (compare Fig. 266 with 
Fig. 259). However, the dark, comma- 
shaped mark inside each opening resem- 
bles that of M. Ventura (compare Fig. 266 
with Fig. 231). As in M. ventura (Fig. 229) 
but unlike all other species in the M. ven- 
tura species group, the embolus tapers 



strongly then curves sharply up and inward 
right at the tip (Fig. 264). The bend in the 
embolus in other M. ventura group species 
is less pronounced (Figs. 236, 257) or not 
so near the tip (Figs. 243, 250). Unlike M. 
ventura, the median apophysis in M. chi- 
lapae is slimmer (Fig. 264) and the sepa- 
ration between the embolus and the basal 
embolic apophysis is relatively greater. 

Variation. Average body length of eight 
females examined 6.5 ixim, range 5.7 to 8.3 
mm. Average body length of three males 
examined 2.9 mm, range 2.3 to 3.5 mm. 

Natural History. Mature specimens 
have been collected in July through Oc- 
tober (Fig. 333). 

Distribution. Southern Mexican states 
between Nayarit and Oaxaca; elevations 
from 1,000 to 2,000 m (Map 13). 

Records Examined. MEXICO Guerrero: 6 mi NE 
Tixtla de Guerrero, 17°39'N, 99°22'W, 16.vii.l984 (J. 
B. Woolley, ADC). Morelos: Cocoyoc, 18°52'N, 
98°59'W, 28.vii.1956 (W. Gertsch & V. Roth, AMNH); 
Cuemavaca, 18°55'N, 99°15'W, 15.X.1944 (N. L. H. 
Krauss, AMNH). Nayarit: 30 mi SE Tepic, 21°12'N, 
104°33'W, 23.xi.1948 (E. S. Ross, CAS); 5 mi NW 
Tepic, 21°32'N, 104°57'W, 13.V.1963 (W. J. Gertsch 
& W. Ivie, AMNH). Oaxaca: 6 mi NE Mitla, 16°59'N, 
96°21'W, 20.viii.l985 (J. Woolley & G. Zolnerowich, 
ADC); Huajuapan, 17°48'N, 97°46'W, l.x.1946 (H. 
Wagner, AMNH); Oaxaca, 17°3'N, 96°43'W, 
17.vii.l955 (C. & R Vaurie, AMNH), 12.viii.l991 (W. 
H. Piel & G. S. Bodner, MCZ); San Bait. Chichica- 
pan, 16°45'N, 96°29'W, 4.viii.l991 (W. H. Piel & G. 
S. Bodner, MCZ). 

Metepeira minima Group 

Female spiders in the M. minima group 
(Metepeira petatlan, Metepeira uninima, 
Metepeira pacifica, Metepeira jamaicensis) 
have a thin scape with epigynal openings 
that are shaped from longitudinal slits 
(Figs. 273, 280, 281) to ovals that are lon- 
ger than wide (Figs. 288, 295). In males, 
the flagellae on the median apophysis are 
set off on a distinctly narrower stalk (Figs. 
271, 278, 286, 293). 

35. Metepeira petatlan new species 
Figures 271-277, 334; IVIap 10 

Holotype. Male from 50 km southeast of Petatlan, 
Guerrero, Mexico, 14.viii.l984, J. B. Woolley, in 



Metepeira • Piel 



81 



MCZ. The specific name is a noun in apposition 
after the locality. 

Description. Female paratype from Pa- 
panoa, Guerrero. Light region around pos- 
terior eye row; carapace darkens posteri- 
orly, then lightens under overhang of ab- 
domen (Fig. 276). Slight rings on patella 
and tibia. Femur I with three macrosetae 
on anterior side, none on anteroventral 
side. Dorsum of abdoinen with typical fo- 
lium, except that white oak leaf pattern 
narrower than most species and set on a 
narrow, remarkably darker brownish dove- 
tail (Fig. 276). Dark lateral band wraps 
around abdomen and stretches up over an- 
terior dorsal portion. Venter with two spots 
on either side of spiracle and slight indi- 
cation of a transverse bar. Wide, white lon- 
gitudinal mark ends at the epigynal groove 
(Fig. 277). Sternum with wide, white lon- 
gitudinal mark v^dening anteriorly (Fig. 
277). Ratio of eye diameters: posterior me- 
dians and anterior medians 1.0, anterior 
laterals 1.3, posterior laterals 1.2. Anterior 
median eyes separated by 1.1 diameters, 
posterior median eyes by 0.7, anterior me- 
dian eyes separated from anterior laterals 
by 1 diameter of anterior lateral eyes, lat- 
eral eyes aliTiost touching. Total length 5 
mm. Carapace 2.2 mm long, 1.7 wide. 
First femur 2.3 mm, patella and tibia 2.5, 
metatarsus 2, tarsus 0.9. Second patella 
and tibia 2 mm, third 1.3, fourth 1.9. 

Male holotype. Carapace uniform 
brown with a lighter median streak just an- 
terior to thoracic depression (Fig. 274). 
Weakly ringed on tibia; distal half of fem- 
ora dark. Feinur I with two macrosetae on 
anterior side. Abdomen is a lighter version 
of the female (Fig. 274). Venter with a 
much reduced and shorter longitudinal 
line, compared to the female (Fig. 275). 
Sternum as in female. Ratio of eye diam- 
eters: posterior medians and anterior me- 
dians 1.0, anterior laterals 1.4, posterior 
laterals 1.4. Anterior median eyes separat- 
ed by 1.1 diameters, posterior median eyes 
by 0.7, anterior median eyes separated 
from anterior laterals by 1.1 diameters of 



anterior lateral eyes, lateral eyes separated 
by 0.2 their diameters. Total length 2 mm. 
Carapace 1 mm long, 0.7 wide. First fe- 
mur 1.2 inm, patella and tibia 1.0, meta- 
tarsus 0.8, tarsus 0.5. Second patella and 
tibia 0.9 mm, third 0.5, fourth 0.8. 

Diagnosis. The female's dorsum can be 
distinguished from other species by the 
unusually dark brown color of the dovetail- 
shaped mark (Fig. 276). Both this mark 
and the white oak leaf pattern have nar- 
rower margins that are more parallel and 
not as wedge-shaped as they are in other 
species (Fig. 276). The epigynum resem- 
bles that of M. uninima by the narrow 
scape and the parallel slitlike openings on 
either side. These openings differ from 
those of M. minima because they have 
conspicuous sclerotized spheres just ante- 
rior to the lateral edge of the slits (Fig. 
273) but which are much farther away in 
M. minima (Fig. 280). The male is unusu- 
ally small and its palp is unique and easily 
distinguished from other species. The lon- 
ger flagellum on the median apophysis is 
needlelike over its entire length, not grad- 
ually tapering (compare Fig. 271 with Fig. 
278). In addition, the long needlelike fla- 
gelluni has a sharper elbow. At the elbow 
it projects away from the palp for a short 
distance and then strongly curves around 
the palp (Fig. 271). 

Natural History. The female paratype 
was found at eye level on a hot, dry, wood- 
ed hillside overlooking the Pacific Ocean 
at an altitude of about 200 m. Her web 
differed considerably from the usual Me- 
tepeira web: instead of an elaborate barrier 
web, it consisted of a much reduced Y- 
shaped structure with a curled leaf in the 
center. The leaf served as a retreat for the 
spider and protected and hid three egg 
sacs. The retreat was suspended very near 
the hub, as opposed to farther away and 
above it, as found in most other species. 
Mature specimens have been collected in 
August through October (Fig. 334). 

Distribution. Western Mexico: Guerrero 
and Sinaloa (Map 10). 



82 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Records Examined. MEXICO Guerrero: 32 mi SE 
Petatlan, 17°14'N, 100°53'W, 14.viii.l984 (J. B. Wool- 
ley, MCZ); Microondas Tamarindos, S. Papanoa, 
17°17'32"N, 101°2'40"W, 18.x. 1994 (W. H. Piel, 
MCZ). Sinaloa: 2 mi S Elota, 23°55'N, 106°48'W, 
ll.Lx.1966 (Jean & Wilton Ivie, AMNH). 

36. Metepeira minima Gertsch 
Figures 278-285, 328; Map 15 

Metepeira minima Gertsch, 1936: 10, fig. 31, 6 . Male 
hoiotype from Edinburg, Texas, in the AMNH, ex- 
amined. Roewer, 1942: 869. Chamberlin and Ivie, 
1942: 67, fig. 174, 9. Levi, 1977: 206, fig. 70-73, 
76-77, 9 ; 74-75, 3 . Bonnet, 1957: 2822. Brignoli, 
1983: 275. 

Description. Female from Celestun, Yu- 
catan, Mexico. Carapace dark brown with 
light region surrounding the eyes, some- 
times extending posteriorly behind the lat- 
eral eyes. Faint pair of darker feather- 
shaped patches in center of carapace (Fig. 
284). Legs same color as carapace; darker 
on distal ends of articles. Femur I with 
row of four macrosetae on anterior side; 
rarely any on anteroventral side. Dorsum 
of abdomen with usual Metepeira folium 
(Fig. 284); venter wide, with long white 
median line surrounded by faint, thin, 
white U-shaped line (Fig. 285). Pair of 
large white spots on either side of spiracle. 
Sternum black with wide, white, uneven 
median line (Fig. 285). Ratio of eye di- 
ameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.2, posterior 
laterals 1.1. Anterior median eyes separat- 
ed by 1.2 diameters, posterior median eyes 
by 0.9, anterior median eyes separated 
from anterior laterals by 1.9 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.2 their diameters. Total length 6.5 
mm. Carapace 2.7 mm long, 2 wide. First 
femur 2.8 mm, patella and tibia 2.8, meta- 
tarsus 2.5, tarsus 0.9. Second patella and 
tibia 2.4 mm, tliird 1.5, fourth 2. 

Male from Celestun, Yucatan, Mexico. 
Carapace dark with light mark anterior to 
thoracic furrow (Fig. 282). Femur I with 
row of four macrosetae on anterior side; 
row of two to five macrosetae on anter- 
oventral side. Distal two-thirds same color 
as carapace, proximal third white. Folium 



of abdomen as in female, except posterior 
half darker than anterior half (Fig. 282). 
Venter of abdomen and sternum as in fe- 
male, except median white line of sternum 
often broken (Fig. 283). Ratio of eye di- 
ameters: posterior medians and anterior 
medians 1.1, anterior laterals 1.4, posterior 
laterals 1.1. Anterior median eyes separat- 
ed by 1.2 diameters, posterior median eyes 
by 0.8, anterior median eyes separated 
from anterior laterals by 1.5 diameters of 
anterior lateral eyes, lateral eyes separated 
by 0.1 their diameters. Total length 3 mm. 
Carapace 1.5 mm long, 1.2 wide. First fe- 
mur 2 mm, patella and tibia 1.9, metatar- 
sus 1.8, tarsus 0.8. Second patella and tibia 
1.6 mm, third 0.8, fourth 1.3. 

Diagnosis. Unlike other members of the 
M. niininia species group, the openings to 
the epigynum of M. minima are narrow 
slits around a scape of variable size (Figs. 
280, 281). The openings of M. petatlan are 
only slightly wider than M. minima, but 
the internal darker sclerotized spheres in 
M. minima sit much farther apart (com- 
pare Fig. 280 with 273). Unlike M. pacifica 
(Fig. 286) and M. jamaicensis (Fig. 293), 
the bent embolus on M. minima tapers 
strongly and is therefore not like a needle 
(Fig. 278). Unlike M. petatlan, the longer 
flagellum on M. minima tapers (Fig. 278) 
and is not thin over its entire length (Fig. 
271). 

Variation. Average body length of 13 fe- 
males examined 5.4 mm, range 3.7 to 6.5 
mm. Average body length of 17 males ex- 
amined 3.4 mm, range 2.5 to 4.5 mm. The 
epigyna of females from the Yucatan pen- 
insula differ noticeably from all others, but 
the males hardly show any distinguishing 
features. These differences can be seen in 
the much wider scape and the greater sep- 
aration between openings in a specimen 
from the Yucatan (Fig. 281), compared to 
a specimen from Tamaulipas (Fig. 280) or 
one from Texas (Levi, 1977: 209, fig. 71). 
In addition, the posterior view of the epi- 
gynum from a Yucatan specimen shows 
the lobes converging to form a V-shape 
(Fig. 279), whereas females outside of the 



Metepeira • Piel 



83 




278 



minima 
(36) 




s 291 




292 




289 ^ 290 



pacifica 

(37) 




298 




297 



jamaicensis 
(38) 



Figures 278-285. Metepeira minima Gertsch (sp, 36 [278,279,281-285] 20°56'N, 90°21'W; [280] 22°30'N, 99°4'W). 278, male 

palpus, mesal. 279, epigynum, posterior. 280,281, epigynum, ventral. 282, male, dorsal. 283, male, ventral. 284, female, dorsal. 

285, female, ventral. 

Figures 286-292. Metepeira pacifica new species (sp. 37; 10°27'N, 85°9'W). 286, male palpus, mesal. 287, epigynum, posterior. 

288, epigynum, ventral. 289, male, dorsal. 290, male, ventral. 291, female, dorsal. 292, female, ventral. 

Figures 293-299. Metepeira jamaicensis Archer (sp. 38; 18°17'N, 76°48'W). 293, male palpus, mesal. 294, epigynum, posterior. 

295, epigynum, ventral. 296, male, dorsal. 297, male, ventral. 298, female, dorsal. 299, female, ventral. 

Scale bars: dorsum and venter figures 1 .0 mm. 



84 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



Yucatan show largely parallel lobes (Levi, 
1977: 209, fig. 71). 

Natural History. Adults have been ob- 
served on tree trunks and on bushes at and 
above 150 cm. They can be found in shad- 
ed areas, which is unusual for Mexican 
Metepeira. Most mature specimens have 
been collected in May through September 
(Fig. 328). Elevations range from sea level 
to just under 2,000 m. 

Distribution. Southern Texas to Hon- 
duras (Map 15). 

Records Examined. HONDUFIAS Copdn: 14°50'N, 
89°9'W, 7.iii.l939 (AMNH); Ruinas, 14°50'N, 
89°9'W, 7.iii.l939 (AMNH). MEXICO Campeche: 
Becan, 18°33'N, 89°30'W, 31.vii.l991 (W. H. Piel & 
G. S. Bodner, MCZ); Campeche, 19°51'N, 90°32'W, 
14.vii.l948 (C. J. Goodnight, AMNH); Champoton 
beach, 19°21'N, 90°43'W, 22.vii.1991 (W. H. Piel & 
G. S. Bodner, MCZ); Seybaplaya, 19°39'N, 90°40'W, 
2.vlii.l949 (C. J. Goodnight, AMNH). Chiapas: 16 mi 
W Cintalpa on rt 190. 16°36'N, 93°51'W, 15.vi.l982 
(F. Coyle, MCZ). Nuevo Leon: 20 km E Montemo- 
relos, Gamine Q. Rayones, 25°16'N, 99°41'W, 
22.vi.1981 (L. Stange, FSCA); Los Cristales, 25°33'N, 
100°12'W, 15.viii.l972 (A. F. Archer, AMNH); Villa 
de Santiago, Hacienda Vista Hermosa, 25°25'N, 
100°9'W, 19.vi.l940 (H. Hoogstraal, MCZ); Villa San- 
tiago, Horsetail Falls, 25°25'N, 100°9'W, 19.vi.l940 
(H. Hoogstraal, MCZ). San Luis Potosi: 20 km W Cd. 
Valles, 21°58'N, 99°11'W, 18.iii.l972 (J. A. L. Cooke, 
AMNH); Cd. Valles, 21°59'N, 99°1'W, 6.vii.l940 (R 
Ran, MCZ), 19.vli.l956 (W. J. Gertsch & V. Roth, 
AMNH), 15.vii.l959 (L. Steude, AMNH); Cd. Valles, 
Hotel Covadonga, 21°59'N, 99°1'W (L. Steude, 
AMNH); Huichichuyan, 21°25'N, 98°55'W, 19.V.1952 
(M. Cazier, W. Gertsch, & R. Schrammel, AMNH); 
Medina, 24°1'N, 100°24'W, 9.Lx.l956 (A. F. Archer, 
AMNH); N section of San Luis Potosi, 22°13'N, 
100°58'W, 8.ix.l956 (A. F. Archer, AMNH); Venado 
Arroyo, 22°56'N, 101°5'W [?], 27.vii.1934 (MCZ). So- 
nora: 8 mi W Alamos, 29°13'N, 110°10'W, 
23.viii.1965 (W. J. Gertsch & R. Hastings, AMNH). 
Tamaulipas: 60 km N Cd Valles, 22°30'N, 99°4'W, 
10.viii.l991 (W. H. Piel & G. S. Bodner, MCZ); Cd. 
Victoria, 23°44'N, 99°8'W, 17.V.1952 (M. Cazier, W. 
Gertsch, & R. Schrammel, AMNH); Laredo road 
near Cd. Victoria, 23°44'N, 99°8'W, 20.viii.l947 (C. 
J. & M. Goodnight, AMNH); rlOl 26km S Tula, 
22°49'N, 99°55'W, 8.Lx.l991 (W. H. Piel & G. S. Bod- 
ner, MCZ); Sisal, 15 mi S Cd. Victoria, 23°38'N, 
99°12'W, 22.vii.1966 (Jean & Wilton Ivie, AMNH). 
Veracruz: 15 mi E Panuco, 22°10'N, 98°3'W, 
29.xi.1941 (A. M. & L. I. Davis, AMNH); Plan del 
Rio, 19°6'N, 96°6'W [?], 26.vii.1956 (W. J. Gertsch & 
V. Roth, AMNH). Yucatan: 20 km E Valladolid, 
20°41'N, 88°2'W, 26.vii.1991 (W. H. Piel & G. S. 



Bodner, MCZ); 3 km S San Felipe, 21°32'N, 
88°14'W, 25.vii.1991 (W. H. Piel & G. S. Bodner, 
MCZ); 5 mi E Sisal salt flat, 21°9'N, 90°5'W, 9.i.l984 
(V. & B. Roth, CAS); Balankanche Cave, 2 km E Chi- 
chen Itza, 20°40'N, 88°33'W, 19.vii.l983 (W. Mad- 
dison, MCZ); beach north of Celestun, 20°56'N, 
90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner, 
MCZ); Chichen Itza, 20°40'N, 88°34'W (C. J. Good- 
night, AMNH), 15.vii.l981 (C. Gold, CAS); Cordil- 
leria Mayapan, 20°28'N, 89°11'W, 6.Lx.l952 (J. & D. 
Pallister, AMNH); Uxmal, 20°22'N, 89°46'W, 
7.ix.l970 (A. F. Archer, AMNH), 23.vii.1991 (W. H. 
Piel & G. S. Bodner, MCZ). USA Texas: 29 mi S 
Sarita, 26°47'N, 97°47'W, 14..xi.l958 (A. Brady, 
MCZ); 1 mi S Pharr, 26°10'N, 98°11'W, 14.xi.l958 
(A. Brady, MCZ); 1 mi S Pharr on U.S. HW 281, 
26°10'N, 98°11'W, 14..xi.l95S (A. Brady, MCZ). 

37. Metepeira pacifica new species 
Figures 286-292, 302; Map 10 

Holotijpe. Male from La Pacifica, Guanacaste, Costa 
Rica, I.ii.l975-2.iii.l975, R. E. Coville, in MCZ. 
The specific name is a noun in apposition after the 
locality. 

Description. Female paratype from La 
Pacifica, Guanacaste, Costa Rica. Carapace 
tan, lighter around eyes. Legs white. Fe- 
mur I with row of three macrosetae on an- 
terior side; none on anteroventral side. 
Dorsum of abdomen white with faint, in- 
distinct folium, darker distally (Fig. 291). 
Gravid females often slightly marbled. 
Venter wide, with long white median line 
surrounded by faint, thin, white U-shaped 
line on black. Pair of large white spots on 
each side of spiracle (Fig. 292). Sternum 
black with wide, white, uneven median 
line (Fig. 292). Ratio of eye diameters: 
posterior medians and anterior medians 
1.0, anterior laterals 1.2, posterior laterals 
1.2. Anterior median eyes separated by 1.2 
diameters, posterior median eyes by 0.9, 
anterior median eyes separated from an- 
terior laterals by 1.8 diameters of anterior 
lateral eyes, lateral eyes separated by 0.2 
their diameters. Total length 4.5 mm. Car- 
apace 2.2 mm long, 1.7 wide. First femur 
2.3 mm, patella and tibia 2.4, metatarsus 
1.9, tarsus 0.8. Second patella and tibia 2 
mm, third 1.2, fourth 1.8. 

Male holotype. Carapace, legs, abdo- 
men as in female, though often darker 
(Figs. 289, 290). Femur I with row of four 



Metepeira • Piel 



85 



300 Dec 

Nov 




301 



gressa, n = 26 



Sep 





Dec 


Jan 




Nov ^ 

< 




z 


'^Mar 


V 




-r* 


W"" 


Aug^ 




\ 






Julyl 


June 





revillagigedo, n = 2 



304 Dec 

Nov 




Apr Sep 




galatheae, n = 304 



tarapaca, n = 23 



July 

rectangula, n = 1 1 



desenderi, n = 46 



308 De': 

Nov 




calamuchita, n = 15 



Nov 


■\ 




\"^ 


OaA 


1 




Am 


'"V 


\y 




V^ 


Aug 


y^ ^ 




May 




July 


rjune 





crassipes, n 




May 
July I June 

nigriventris, n = 44 



Aug NC^ ^ jP\ ^^y 

July I June 

compsa, n = 279 




datona, n = 66 



jamaicensis, n = 24 



glomerabilis, n = 33 



cajabamba, n = 1 1 




Aug "V^ 1 Js/ May 
July June 

roraima, n = 11 



maya, n = 1 3 



Aug ^yl ■" \^ May 

July June 

inca, n = 29 



Aug ^^ ^1 J^ May 
July ' June 

arizonica, n = 45 



Figures 300-319. Circular liistograms depicting relative seasonal abundance of collecting events for mature spiders. 300-308. 
Primarily collected during the northern hemisphere winter and spring seasons. 300, Metepeira gressa; 301, Metepeira revillagi- 
gedo; 302, Metepeira pacifica; 303, Metepeira karkii; 304, Metepeira galatheae; 305, Metepeira tarapaca; 306, Metepeira rec- 
tangula; 307, Metepeira desenderi; 308, Metepeira calamuchita. 309-314. Generally collected throughout. 309, Metepeira cras- 
sipes; 310, Metepeira nigriventris; 311, Metepeira compsa; 312, Metepeira datona; 313, Metepeira jamaicensis; 314, Metepeira 
glomerabilis. 315-319. Primarily collected during the Northern Hemisphere summer season. 315, Metepeira cajabamba; 316, 
Metepeira roraima; 317, Metepeira maya; 318, Metepeira inca; 319, Metepeira arizonica. 



86 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



macrosetae on anterior side; two to three 
on antero ventral side. Median white line 
on sternum often broken (Fig. 290). Ratio 
of eye diameters: posterior medians and 
anterior medians 1.0, anterior laterals 1.6, 
posterior laterals 1.6. Anterior median 
eyes separated by 1.1 diameters, posterior 
median eyes by 0.6, anterior median eyes 
separated from anterior laterals by 1.4 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 0.3 their diameters. To- 
tal length 3.4 mm. Carapace 1.7 mm long, 
1.2 wide. First femur 2.2 mm, patella and 
tibia 2, metatarsus 1.7, tarsus 0.8. Second 
patella and tibia 1.6 mm, third 0.9, fourth 
1.3. 

Diagnosis. Unlike other members of the 
M. minima species group, M. pacifica and 
M. jamaicensis share very light pigmenta- 
tion, and the embolus in both species is 
needle-shaped (Figs. 286, 289, 291, 293, 
296, 298). Unlike M. jamaicensis, the fla- 
gellae on the median apophysis of M. pa- 
cifica are set off on a short, wide stalk (Fig. 
286) rather than a long, thin one (Fig. 
293). A ventral view of the epigynum in 
M. pacifica shows a ridge under the scape 
that is more V-shaped (Fig. 288), com- 
pared to a straighter line (Fig. 295). 

Variation. Average body length of three 
females examined 5.3 mm, range 4.5 to 5.8 
mm. Average body length of five males ex- 
amined 3 mm, range 2.8 to 3.4 mm. Most 
specimens appear whitish, but the loss of 
pigment is variable, especially among 
males. 

Natural History. Specimens have been 
collected from the wasp nests of Trtjpar- 
gilum nitidum, T. tenoctitlan, and T. hen- 
soni. In Costa Rica, mature specimens 
have been collected in November through 
February; Honduras and Nicaragua in July 
(Fig. 302). Altitudes range from 100 to 
2,600 m. 

Distribution. Costa Rica, Honduras, and 
Nicaragua (Map 10). 

Records Examined. COSTA RICA Guanacaste: 4 
km NW Canas, La Pacifica, prey of Trypargilum ben- 
soni, 10°27'N, 85°9'W, 29.1.1975 (R. E. Coville, 
MCZ); 4 km NW Canas, La Pacifica, prey of Trypar- 



gilum nitidum, 10°27'N, 85°9'W, I.ii.l975-2.iii.l975 
(R. E. Coville, MCZ); 4 km NW Caiias, La Pacifica, 
prey of Trypargilum tenoctitlan, 10°27'N, 85°9'W 
l.ii.l975 (R. E. Coville, MCZ); 4 km NW Canas, La 
Pacifica, wasp collected, 10°27'N, 85°9'W, 16.ii.l975 
(R. E. Coville, MCZ); Bagaces, Palo Verde, 10°21'N, 
85°21'W 19.1.1978 (W. Eberhard, MCZ); ca. Canas, 
10°25'N, 85°7'W, 15.xi.l982 (W Eberhard, MCZ); 
Canas, 10°25'N, 85°7'W 15.xl.l982 (W Eberhard, 
MCZ). Puntarena.s: Elnca San Miento-Sialas, 10°9'N, 
84°54'W, 5.11.1976 (Roth & Schroepfer, AMNH). San 
Jose [?]: Santa Maria, 9°39'N, 83°58'W, 15.1.1930 
(Dodge, MCZ). HONDURAS Tegucigalpa: Teguci- 
galpa, 14°6'N, 87°13'W, l.vii.l948 (Clarke, AMNH). 
NICARAGUA Managua: Laguna de Jiloa, SW Ma- 
nagua, campsite, 12°13'N, 86°19'W 8.vil.l970 (S. 
Riechert, SR). 

38. Metepeira jamaicensis Archer 
Figures 293-299, 313; Map 15 

Metepeira jamaicensis Archer, 1958: 16, fig. 33, 9 . 
Female holotype from Port Henderson, St. Cath- 
erine Parish, Jamaica, in the AMNH, examined. 

Metepeira rmnima: — Levi, 1977: 206, 208. BrignoH, 
1983: 275. Erroneous synonymy. 

Description. Female from Saint Mary's 
Parish, Strawberry Fields near Robin s Bay 
and Green Castle, Jamaica. Carapace dirty 
brown, white around eyes, central w^hite 
wedge (Fig. 298). Legs whitish yellow; 
slight rings on legs II and III. Femur I 
with row of three to four macrosetae on 
anterior side; none or only a few very fine 
setae on anteroventral side. Dorsal folium 
lighter than in most species; fleur-de-lis 
white on speckled light gray (Fig. 298). 
Venter brownish gray with white margins. 
Wide median white line with pair of large 
white spots on either side of spiracle (Fig. 
299). Sternum brownish black with wide, 
white line widening anteriorly, sometimes 
broken in center (Fig. 299). Ratio of eye 
diameters: posterior medians and anterior 
medians 1.0, anterior laterals 1.2, posterior 
laterals 1.1. Anterior median eyes separat- 
ed by 1.4 diameters, posterior median eyes 
by 0.7, anterior median eyes separated 
from anterior laterals by 2 diameters of an- 
terior lateral eyes, lateral eyes separated by 
0.2 their diameters. Total length 5.1 mm. 
Carapace 2.1 mm long, 1.5 wide. First fe- 
mur 2.1 mm, patella and tibia 2.2, meta- 



Metepeira • Piel 



87 




Aug NC^ ^t \^ May 
July I June 

Ventura, n = 35 



Aug N4/ 1 J\ May 
July I June 

grandiosa grandiosa, n = 4 



22 


Dec J 


Jan 




Nov , 






Feb 


OctN 






-^Mar 


Sepi^ 






J*, 


Aug 


July 


June 


•'^^May 




gosoga, n = 2 



Aug 7^^ J\ May 

July I June 

incrassata, n = 33 




Aug N/ W\ \; ^^"^ 

July I June 

grandiosa alpina, n = 5 



326 


Dec 


Jan 




Nov 




7^ 


Feb 


Oct /^ 




^ 


Am^ 


Sep\ 




J^ 


Jap. 


Aug 




\ 


May 




July 


June 





327 


Dec 


Jan 




Nov 




7^ 


Feb 


Oct A 




■V 


"^Mar 


Sepi^ 




^ 


/ Apr 


Aug^ 




\ 


May 




July 


June 





celestun, n = 8 



vigilax, n = 10 



28 


Dec J 


Jan 




NOV/ 


\ 




Feb 


Oct?^ 


f^ 




\ Mar 


Sep\> 


V 




\ / ^^' 


Aug^ 


/r 




May 




juijn 


June 





329 



Sep 



Dec 
Nov J\ 


Jan 


^\^Mar 




^ 


J*" 


AugNc^^ 


\ 


May 


J^ 


\lune 





30 

Nov X 


Dec 


Jan 


Feb 


Oc.?^ 


^ 




-^Mar 


Sep^ 


K 




-^Apr 


AugV 


;4 




May 




July 


June 






minima, n = 39 



uncata, n = 6 



pimungan, n = 6 



Comanche, n = 2 



332 


Dec 


Jan 




Nov 




y^ 


^Feb 


OoN 




V 


-A Mar 


.pi^ 




^ 


J Apr 


Aug 




\ 


May 




Julyl 


June 





333 


Dec J 


Jan 




Nov 


\ 




Feb 


Oct /^ 


j^ 




•^A Mar 


Sep\ 


\t 




Ja. 


Aug 


YJ 




May 




July 


Fjune 





34 

Nov X 


Dec 


Jan 


Feb 


o„/V 


J^ 




'^A Mar 


sS^ 


V 




■\ / ^P' 


Aug^ 


^ 




May 




July 


Ijune 





335 


Dec 


Jan 




Nov 


\ 




^Feb 


OctN 


1^ 




-^Mar 


'"U 


|L 




J Apr 


Aug 


^ 




/ 

May 




jiiiTl 


rTune 





lacandon, n = 10 



chilapae, n = 15 



petatlan, n = 3 



spinipes, n = 1 16 



336 


Dec J 


Jan 




Nov , 






Feb 


Oct r^ 






•^A Mar 


Sep y . 

V4 






^Apr 


Aug 


July 


June 


May 



atascadero, n = 1 7 




triangularis, n = 19 



Figures 320-337. Circular histograms depicting relative seasonal abundance of collecting events for mature spiders. 320-331, 
Primarily collected during the northern hemisphere summer season. 320, Metepeira ventura; 321, Metepeira grandiosa gradiosa; 
322, Metepeira gosoga; 323, Metepeira ventura; 324, Metepeira olmec; 325, Metepeira grandiosa alpina; 326, Metepeira celes- 
tun; 327, Metepeira vigilax; 328, Metepeira minima; 329, Metepeira uncata; 330, Metepeira pimungan; 331, Metepeira comanche. 
332-337. Primarily collected during the northern hemisphere fall season. 332, Metepeira lacandon; 333, Metepeira ctiilapae; 
334, Metepeira petatlan; 335, Metepeira spinipes; 336, Metepeira atascadero; 337, Metepeira triangularis. 



88 Bulletin Museum of Comparative Zoology, Vol. 157, No. 



tarsus 1.8, tarsus 0.8. Second patella and 
tibia 1.9 mm, third 1.2, fourth 1.7. 

Male from same locality as female. Male 
carapace, dorsum, venter, sternum darker 
and more contrasty version of female 
(Figs. 296, 297). All legs ringed. Femur I 
with row of three macrosetae on anterior 
side; none on anteroventral side. Ratio of 
eye diameters: posterior medians and an- 
terior medians 0.9, anterior laterals 1.2, 
posterior laterals 1.2. Anterior median 
eyes separated by 1.3 diameters, posterior 
median eyes by 0.9, anterior median eyes 
separated from anterior laterals by 1.2 di- 
ameters of anterior lateral eyes, lateral 
eyes separated by 2.3 their diameters. To- 
tal length 2.3 mm. Carapace 1.2 mm long, 
0.9 wide. First femur 1.4 mm, patella and 
tibia 1.3, metatarsus 1.0, tarsus 0.5. Sec- 
ond patella and tibia 1.2 mm, third 0.6, 
fourth 0.9. 

Diagnosis. Unlike other members of the 
M. minima species group, M. jamaicensis 
and M. pacifica share very light pigmen- 
tation and the embolus in both species is 
needle-shaped (Figs. 286, 289, 291, 293, 
296, 298). Unlike M. pacifica, the flagellae 
on the median apophysis of M. jamaicensis 
are set off on a long, thin stalk (Fig. 293) 
rather than a short, wide one (Fig. 286). A 
ventral view of the epigynum in M. jamai- 
censis shows a ridge under the scape that 
almost forms a straight line (Fig. 295), 
compared to a V-shape (Fig. 288). 

Variation. Average body length of elev- 
en females examined 5 mm, range 4.2 to 
6.1 mm. Average body length of four 
males examined 2.4 mm, range 2.3 to 2.6 
mm. 

Natural History. Mature specimens 
have been collected in July through March 
(Fig. 313). 

Distribution. Primarily in Jamaica and 
Haiti (Map 15), near sea level. 

Records Examined. BRITISH WEST INDIES 

Grand Cayman Island: 19°20'N, 81°10'W, 15.ii.l960 
(R. A. Lewin, MCZ). HAITI Departement de 
L'Ouest: Port-au-Prince, 18°32'N, 72°20'W, 
19.vii.l955 (A. F. Archer, AMNH), 20.vii.l955 (A. F. 
Archer, AMNH). Dept. de L'Artibonite: Saint-Marc, 



19°7'N, 72°42'W, 15,i.l913 (W. M. Mann, MCZ). JA- 
MAICA Cornwall: Montego Bay, 18°28'N, 77°55'W, 
l.iii.l984 (L. E. Schulten Jr, MCZ). Middlesex: 3 mi 
E Old Harbor, 17°56'N, 77°10'W, 21.X.1957 (A. M. 
Chickering, MCZ); Christiana, 18°10'N, 77°29'W, 
13.xi.l957 (A. M. Chickering, MCZ), 15.vii.l960 (C. 
& P Vaurie, AMNH), 17.vii.l960 (C. & P Vaurie, 
AMNH); Strawberry Fields near Robin's Bay and 
Green Casde, 18°17'N, 76°48'W, 23.iii.1972 (H. W., 
L. & F Levi, MCZ), 25.iii.1972 (H. W., L. & F. Levi, 
MCZ), 26.iii.1972 (H. W., L. & F Levi, MCZ). Saint 
Ann: Roaring River, 18°24'N, 77°9'W [?], 8.ii.l946 
(B. Heineman, AMNH); Saint Ann's Bay, 18°26'N, 
77°8"W, 20.xi.l959 (A. M. Nadler, AMNH). Saint 
Catherine: E Green Harbour, S slope of Healdishire, 
17°53'N, 76°51"W, 12.viii.l958 (A. F. Archer, 
AMNH). St. Andrews: Ferry, 9/10 mi on Spanishtown 
Road, 18°2'N, 76°53"W, 26.vii.1955 (A. F Archer, 
AMNH). Surrey: Kingston, Mona Road, pasture, 
17°59'N, 76°24"W, lO.x.1957 (A. M. Chickering, 
MCZ); Roselle Falls, 24 mi E Kingston, 17°59'N, 
76°24"W, 29.X.1957 (A. M. Chickering, MCZ); Saint 
Andrew, 18°4'N, 76°45'W, 7.X.1957 (A. M. Chicker- 
ing, MCZ). Trelawny: Falmouth, 18°30'N, 77°39"W, 
20.vii.l960 (C. & P. Vaurie, AMNH). Westinorland: 
Negril, 18°16'N, 78°21'W, 24.iii.1955 (A. M. Nadler, 
AMNH); 'Whitehouse, 18°4'N, 77°58"W, 26.iii.1955 
(A. M. Nadler, AMNH). 

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INDEX 

Valid taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer to 
species descriptions. 



acostai, Metepeira 63 

Aculepeira 8 

alpina, Metepeira 23, 24 

Amazonepeira 8 

Arachnidornyia 10 

Araneus 8 

arizonica, Metepeira 9, 19, 65, 66 

atascadero, Metepeira 9, 16, 18, 67, 69 

bani, Metepeira 63 

biogeography 9, 10 

cajabamba, Metepeira 9, 12, 17, 26, 31 

calamuchita, Metepeira 14, 18, 39, 42 

cancriforrnis, Gasteracantha 10 

celestun, Metepeira 9, 10, 16, 19, 74, 75 

cereicola, Metepeira 43 

chilapae, Metepeira 9, 16, 19, 78, 79 

chilapica, Metepeira 78 

Comanche, Metepeira 14, 19, 60, 61 

compsa group, Metepeira 47 

compsa, Aranea 48 

compsa, Metepeira 7, 9, 10, 11, 14, 17, 48, 49 

crassipes, Metepeira 7, 9, 10, 16, 19, 77, 79 

Cyrtophora 8, 9 

dakota, Metepeira 23, 24 

daytona, Metepeira 6, 7, 8, 9, 10, 12, 17, 20 

desenderi, Metepeira 6, 10, 12, 17, 19, 21, 25 

digital photography 4 

dominicana, Metepeira 30 

dorsal folium 6 

douglasi, Metepeira 34 

ensenada, Metepeira 71 



epigynum 8 

foxi group, Metepeira 19 
foxi, Metepeira 8, 9, 10, 12, 17 
galatheae, Araneus 43 
galatheae, Epeira 43 
galatheae, Metepeira 9, 14, 18, 43, 45 
glomerabilis, Araneus 28 
glomerabilis, Epeira 28 
glomerabilis, Metazygia 28 
glomerabilis, Metepeira 9, 12, 17, 28, 31 
gosoga, Metepeira 14, 18, 59, 61 
grandiosa alpina, Metepeira 9, 12, 17, 24, 25 
grandiosa grandiosa, Metepeira 9, 10, 12, 17, 23, 25 
grandiosa palustris, Metepeira 9, 12, 17 
grandiosa, Metepeira 23 
gressa, Epeira 54 
gressa, Metazygia 54 
gressa, Metepeira 9, 18, 54, 57 
gressus, Araneus 54 
grinnelli, Metepeira 34 
habitats 9 

inca, Metepeira 6, 14, 17, 18, 57, 58 
incrassata group, Metepeira 55 
incrassata, Metepeira 8, 9, 10, 11, 16, 19, 68, 69 
inerma, Metepeira 20 
jamaicensis, Metepeira 9, 17, 19, 83, 86 
josepha, Metepeira 77 
Kaira 6, 8 
karkii, Araneus 46 
karkii, Metepeira 9, 13, 18, 45, 46 
koepckeonim, Araneus 6 



92 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1 



labyrinthea grinnelli, Aranea 34 

labyrinthea grinnelli, Epeira 34 

labyiinthea grinnelli, Metepeira 34 

labyrintliea group, Metepeira 33 

labyrinthea, Epeira 21 

labtjrinthea, Metepeira 7, 9, 10, 14, 17, 32, 34, 46, 48 

labyrintheus, Araneus 48 

lacandon, Metepeira 9, 14, 17, 35, 37 

latigyna, Metepeira 48 

lindae, Arachnidoijia 10 

maija, Metepeira 9, 14, 18, 56, 57 

measurements 5 

Mecynogea 8, 9 

median apophysis 8 

Metepeira 5 

minima group, Metepeira 80 

minima, Metepeira 9, 10, 12, 16, 19, 81, 83, 86 

nigriventris group, Metepeira 38 

nigriventris, Araneus 38 

nigriventris, Epeira 38 

nigriventris, Metepeira 9, 10, 11, 12, 18, 38, 39 

ocosingo, Mecynogea 10 

olmec, Metepeira 9, 16, 19, 5.9, 61 

pacifica, Metepeira 16, 19, 83, 84 

palomara, Metepeira 23 

palp 8 

palustris, Metepeira 23 

perezi, Metepeira 48 

petatlan, Metepeira 16, 19, 79, 80 

pimungan, Metepeira 8, 9, 16, 19, 62, 65 

predation 10 

rayorae, Arachnidomyia 10 

rectangula, Epeira 32 



rectangula, Metepeira 5, 6, 9, 12, 17, 32, 35 

rectangulata, Metepeira 32 

revillagigedo, Metepeira 16, 19, 73, 75 

roraima, Metepeira 9, 12, 18, 49, 53 

Salei, Epeira 68 

salei, Metepeira 70 

sallei, Aranea 70 

sallei, Araneus 70 

santa, Aranea 28 

scitulus, Araneus 54 

seditiosa, Epeira 54 

seditiosa, Eustala 54 

seditiosa, Metepeira 54 

seditiosus, Araneus 54 

Singa 6 

species groups 10 

spinipes, Araneus 34 

spinipes, Metepeira 5, 6, 8, 9, 14, 17, 34, 35 

suspended retreat 8 

tarapaca, Metepeira 7, 9, 14, 18, 39, 40 

triangularis, Metepeira 9, 15, 18, 63, 65 

Trypargilum 10 

uncata, Metepeira 9, 16, 19, 75, 76 

uncatus, Araneus 76 

vaurieomm, Metepeira 48 

Ventura group, Metepeira 71 

Ventura, Metepeira 10, 16, 19, 69, 71 

vigilax group, Metepeira 26 

vigilax, Araneus 30 

vigilax, Epeira 29 

vigilax, Metepeira 9, 10, 12, 17, 26, 30, 31 

virginensis, Metepeira 48 

web 8 

Zygiella 6 



OF THE 



seum 



(US ISSN 0027-4100) 



Type Specimens of Recent Mammals 
in the Museum of Comparative Zoology 



K. M. HELGEN ANDT. L. McFADDEN 



1V1CZ 
LIBRARY 

JUL 3 2001 

HA^VARO 
y N L V ii:^ R -^ ^"^ ' 



HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 157, NUMBER 2 
8 JUNE 2001 



(US ISSN 0027-4100) 



PUBLICATIONS ISSUED 

OR DISTRIBUTED BY THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

HARVARD UNIVERSITY 



Breviora 1952- 

bulletin 1863- 

Memoirs 1865-1938 

JoHNSONiA, Department of Mollusks, 1941-1974 

Occasional Papers on Mollusks, 1945- 

SPECIAL PUBLICATIONS. 

1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and 
Evolution of Crustacea. 192 pp. 

2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- 
dinidea (Mollusca: Bivalvia). 265 pp. 

3. Sprinkle, J-., 1973. Morphology and Evolution of Blastozoan Echino- 
derms. 284 pp. 

4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the 
Present Day. 236 pp. 

5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology 
and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 

725 pp. 

6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. 

Other Publications. 

Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. 
Reprinted 1964. 

Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of 
Insects. {Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971. 

Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. 

Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- 
ternational Symposium on Natural Mammalian Hibernation. {Bulletin 
of the M. C. Z, Vol 124.) 

Orinthological Gazetteers of the Neotropics (1975-). 

Peter's Check-list of Birds of the World, vols. 1-16. 

Proceedings of the New England Zoological Club 1899-1947. (Complete 
sets only.) 

Proceedings of the Boston Society of Natural History. 

Price list and catalog of MCZ publications may be obtained from Publica- 
tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, 
Massachusetts 02138, U.S.A. 

This publication ha': been printed on acid-free permanent paper stock. 

©The President and Fellows of Harvard College 2001. 



TYPE SPECIMENS OF RECENT MAMMALS IN THE MUSEUM OF 
COMPARATIVE ZOOLOGY 

K. M. HELGEN AND T. L McFADDEN^ 

CONTENTS 



Abstract - 94 

Introduction 94 

Authors of Type Descriptions 94 

Definitions and Organization 96 

Taxa Included in This Catalogue .._ 97 

Abbreviations 98 

Acknowledgments 98 

Paralectotype Series 98 

Accounts of Type Specimens 99 

Order Didelphimoi"phia 99 

Family Caluromyidae 99 

Family Marmosidae 99 

Family Didelphidae 100 

Order Dasyuromoqohia 100 

Family Dasyuridae 100 

Order Peramelia 100 

Family Peroiyctidae 100 

Order Cingulata 101 

Family Dasypodidae 101 

Order Afrosoricida 101 

Family Chiysochloridae 101 

Family Tenrecidae 101 

Order Rodentia 103 

Family Aplodontidae 103 

Family Sciuridae 103 

Family Castoridae 109 

Family Geomyidae 110 

Family Heteromyidae 111 

Family Dipodidae 112 

Family Muridae 112 

Subfamily AiAdcolinae 112 

Subfamily Cricetinae 117 

Subfamily Cricetomyinae 117 

Subfamily Dendromurinae 117 

Subfamily Gerbillinae 117 

Subfamily Murinae 117 

Subfamily Nesomyinae 120 

Subfamily Otomyinae 120 

Subfamily Sigmodontinae 121 

Family Pedetidae 128 



^ Mammal Department, Museum of Comparative 
Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 



Family Myoxidae 129 

Family Bathyergidae 129 

Family Erithizontidae 130 

Family Dasyproctidae 130 

Family Agoutidae 130 

Family Octodontidae 130 

Family Echimyidae 131 

Family Capromyidae 131 

Order Lagomoqiha 133 

Family Ochotonidae 133 

Family Leporidae 133 

Order Scandentia 134 

Family Tupaiidae 134 

Order Primates 135 

Family Indridae 135 

Family Daubentoniidae 135 

Family Galagonidae 135 

Family Cebidae 135 

Family Hylobatidae 136 

Family Hominidae 136 

Order Lipotyphla 136 

Family Nesophontidae 136 

Family Solenodontidae 137 

Family Soricidae 137 

Family Talpidae 140 

Order Chiroptera 140 

Family Pteropodidae 140 

Family Emballonuridae 142 

Family Nycteridae 142 

Family Rhinolophidae 142 

Family Mormoopidae 143 

Family Phyllostomidae 143 

Family Molossidae 145 

Family Vespertilionidae 145 

Family Thyi^opteridae 147 

Order Artiodactyla 147 

Family Tayassuidae 147 

Family Monodontidae 148 

Family Phocoenidae 148 

Family Cervidae 148 

Family Bovidae 149 

Order Carnivora 149 

Family Canidae 149 

Family Ursidae 151 



Bull. Mus. Comp. ZooL, 157(2): 93-181, June, 2001 93 



94 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Family Procyonidae 151 

Family Mustelidae 152 

Family Mephitidae 154 

Family Viverridae 155 

Family Heipestidae 155 

Family Felidae 155 

Order Cimolesta 156 

Family Manidae 156 

References 157 

Index 171 

Abstract. The Mammal Department at the Mu- 
seum of Comparative Zoology houses name-bearing 
types of 342 species-group taxa of Recent inammals. 
The type collection consists of 327 holotypes, 2 lec- 
totypes, 4 complete syntype series, and 9 partial syn- 
type series. This catalogue notes information on the 
type locality, collector, date of collection, present con- 
dition, original publication, and synonyms for all 
name-bearing types in the Mammal Department. 
Comments on the taxonomic and historical impor- 
tance of many type specimens are included. Lecto- 
types for a number of taxa are designated for pur- 
poses of taxonomic consistency. 

INTRODUCTION 

"Some of them are more or less historic 
specimens," wrote the great mainiTialogist 
Glover M. Allen in 1931, "whose location 
has undoubtedly in many cases been lost 
sight of, so that it may be of value to make 
the present record" (1931: 230). Allen, 
then Curator of Mammals at the Museum 
of Comparative Zoology at Harvard Uni- 
versity, was referring to the type speci- 
mens of inammals in the museum, the 
subject of a listing he published that year. 
Allen's words resound with even greater 
truth today than when they were written 
70 years ago. Since that time, the inuse- 
um s holdings of mainmal type specimens 
have grown considerably. Furthermore, 
type specimens of ixiamiTials housed in the 
MCZ have sometimes been credited to the 
collections of other institutions or consid- 
ered to no longer exist, making the need 
for a new catalogue obvious. 

Type specimens are biological reference 
points that lend objectivity to taxonomy 
and are thus of critical importance in sys- 
teinatic investigations; accordingly, "eveiy 
institution in which name-bearing types 
are deposited should publish lists of name- 
bearing types in its possession or custody" 



(International Commission on Zoological 
NoiTienclature, 1999: 79). Our duty in this 
regard is thus long overdue, and this cat- 
alogue should sei've to fulfill that obliga- 
tion. Without doubt, it will alleviate con- 
fusion in the scientific coiximunity as to the 
w^hereabouts of certain specimens, thought 
to be lost, especially those type speciinens 
that were acquired from Guillaume Gran- 
didier. In addition, it should bring to light 
information, fonnerly unavailable, on this 
museum s veiy notable holdings of name- 
bearing types. 

All taxonoinic judgments in this work, 
including new name combinations and lec- 
totype designations, reflect the decision of 
the first author (KMH) alone and should 
be cited accordingly. 

AUTHORS OF TYPE DESCRIPTIONS 

Four maminalogists authored the over- 
whelming majority of names based on type 
specimens in the MCZ: Glover Allen, Out- 
ram Bangs, Guillaume Grandidier, and 
Barbara Lawrence. Following are brief bi- 
ographies of these four outstanding mam- 
malogists. 

Glover M. Allen (1879-1942) | 

Curator of Mammals at the MCZ from 
1924 until his death in 1942, Glover Allen 
began his work in the Mammal Depart- 
ment in 1907. He was known as a careful, 
dependable researcher and an outstand- 
ing, patient teacher. His goal in all en- 
deavors was to increase the sum total of 
knowledge about the mammals of the 
world. Although much of his career was 
spent in the museum studying, as he put 
it, "the dried remains of animals" (Barbour 
et al, 1943: 300), he also traveled widely 
throughout the world collecting specimens 
as well as observing and learning about liv- 
ing animals. He once commented that the 
actual knowledge of living creatures could 
all too often be summed up by saying 
"when we found it, it ran like hell, where- 
upon we shot it!" (Barbour et al., 1943: 
300). He wrote prolifically; his bibliogra- 
phy of publications is 81 pages long, the 



Type Specimens of Recent Mammals • Helgen and McFadden 95 



first of which he pubHshed when he was 
only 11 years old (Lawrence 1947a: 1). 
Holotypes for 96 taxa, which Allen de- 
scribed alone or with colleagues, are de- 
posited in the MCZ. 

Outram Bangs (1863-1932) 

Outram Bangs, Curator of Mammals 
from 1899 to 1924, "was one of those for- 
tunate mortals, born with a love of nature 
and the outdoors which rule their entire 
lives" (Peters 1933: 265). Bangs authored 
135 MCZ mammal names (only one co- 
authored!). He and brother Edward began 
collecting as boys using slingshots and 
horsehair nooses. He was an early ecolo- 
gist, saving Microtus breweri by killing the 
feral cat population on Massachusetts' 
Muskeget Island and restocking the mouse 
from a tiny islet across the channel. Al- 
though his greatest passion was the natural 
history of birds, he was also fascinated with 
mammals and served as curator of both 
departments in the MCZ. He decided to 
systematically collect the mammals of east- 
ern North America in about 1890. He be- 
gan by trapping in New England and later 
made trips to the southeastern United 
States and Canada. Other collectors assist- 
ed Bangs, expanding the collection area to 
western North America and south to Cen- 
tral America. His precision and organiza- 
tion were legendaiy, and today his mam- 
mal collection of more than 10,000 speci- 
mens, donated to the Museum in 1899, re- 
mains one of the best curated and most 
informative in the department. A complete 
list of Bangs' scientific publications was 
compiled by Porter (1943). 

Guillaume Grandidier (1873-1957) 

French explorer and scientist Guillaume 
Grandidier, son of naturalist Alfred Gran- 
didier, authored and coauthored descrip- 
tions of 13 taxa whose types were donated 
to the MCZ in 1947, along with his exten- 
sive personal collection of Malagasy mam- 
mals. The collection was purchased by 
Robert Barbour and donated to the MCZ 
in honor of his brother, MCZ director 



Thomas Barbour. Between 1898 and 1902, 
Grandidier explored the center and south- 
ern portions of Madagascar, collecting 
specimens and describing the geography 
of the area. Through his writings he 
brought the unique fauna of this remote 
region to the attention of the scientific 
world. He was known for his devotion to 
the careful acquisition of knowledge, 
whether it was geographical, historical, or 
scientific. His work was honored by both 
the scientific community and the French 
government (Chapus, 1953). 

Barbara Lawrence (1909-97) 

After graduating from Vassar College in 
1931, Barbara Lawrence became a volun- 
teer at the MCZ. She was encouraged by 
Dr. Glover Allen to do her own research, 
and in the late 1930s she made field trips 
to the Philippines and Sumatra to collect 
mammals. In 1952 she was appointed Cu- 
rator of Mammals, a position she held un- 
til her retirement in 1976. Her areas of 
scientific interest were many, ranging from 
echolocation in w^hales to zooarchaeology, 
as well as the more traditional mammalog- 
ical pursuit of taxonomy. "She once wrote 
'. . . to know and love a bit of the world 
so well that you can give it to someone else 
... is a rare talent' " (Rutzmoser, 1999: 
1049), certainly a talent Barbara Lawrence 
had in abundance. She authored or coau- 
thored 20 MCZ names. 

The specimens described by these au- 
thors and others are the result of world- 
wide collecting by numerous expeditions 
and individuals, listed in the accounts that 
follow. A handful of type specimens in the 
MCZ were formerly in the collection of 
the National Museum of Natural Histoiy 
in Washington, D.C. According to Glover 
Allen, these were obtained via exchanges 
"at a time when 'duplicates' were more 
freely disposed of, [and would] prove to be 
cotypes or even actual types" (1931: 229). 
The collections of the Boston Society of 
Natural Histoiy, once contained in the 
Boston Museum of Science, were trans- 



96 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



ferred to the Museum of Comparative Zo- 
ology over the course of the 20th century. 
Several types are among these specimens, 
which consist primarily of mammals from 
the New England region. 

DEFINITIONS AND ORGANIZATION 
Type Categories 

Type specimens are categorized as one 
of the following: 

Holotype. The single specimen desig- 
nated as name-bearer in the original pub- 
lished description of the species-group tax- 
on (Article 73.1, International Commission 
on Zoological Nomenclature, 1999: 79). 

Syntype. One of multiple specimens on 
which a species-group name is equally 
based, when no holotype is specified in the 
original description and no subsequent 
designation of a lectotype has been pub- 
lished (Article 73.2, International Com- 
mission on Zoological Nomenclature, 
1999: 81). 

Lectotype. One of multiple specimens 
upon which a species-group name is orig- 
inally based, designated in a publication 
subsequent to the original description to 
become the unique name-bearer (Article 
74, International Commission on Zoologi- 
cal Nomenclature, 1999: 82). 

Neotype. A specimen chosen as the 
name-bearer of a species-group taxon, "if 
no holotype, lectotype, syntype, or prior 
neotype is believed to exist" (Article 75, 
International Commission on Zoological 
Nomenclature, 1999: 84). There are no 
neotypes in the MCZ mainmal collection. 

Paratype. A specimen other than the 
holotype (if designated originally) that is 
mentioned in the original description of a 
species-group taxon. Paratypes of species- 
group taxa whose holotypes are housed in 
other institutions are not mentioned in this 
catalogue, although inany such specimens 
exist in this museum. 

Paralectotype. If no holotype is desig- 
nated, a specimen other than the lectotype 
(if designated subsequently) that is men- 
tioned in the original description of a spe- 



cies-group taxon. Paralectotypes of spe- 
cies-group taxa whose lectotypes are 
housed in other institutions are discussed 
below, before the accounts for name-bear- 
ing types. 

Locality 

This categoiy includes country, second- 
level geopolitical division (state, depart- 
ment, territory, province, or district), col- 
lection site, and altitude where available. 
The geographic name given for a type lo- 
cality is that found in the original publi- 
cation. When the description of the col- 
lection locality does not include current 
geopolitical divisions, that information is 
provided in parentheses. If an original 
name is no longer used, an equals sign ( = ) 
is included within the parentheses to des- 
ignate an equivalent modern name. Where 
altitude was originally given in feet, it has 
been converted to meters and included in 
parentheses. 

Sources used for the current names are 
from the most current available Gazetteers 
of the United States Board on Geographic 
Names, the 10th comprehensive edition of 
the Times Atlas of the World, the Colum- 
bia Gazetteer of the World, and the Or- 
nithological Gazetteers of South America. 

Account Organization 

The type locality, collector, date of col- 
lection, and present condition of each 
specimen are noted. The publication of 
the original description is cited for each 
specimen. Many names have changed in 
rank or synonymy since their origin; in 
these cases, the name by which a taxon is 
known today is noted, with a citation of the 
publication in which that name combina- 
tion was first employed for that taxon. 
Comments are offered for most entries to 
provide additional information or to dispel 
potential sources of confusion. 

The format of this catalogue is largely 
borrowed from the most recent type cat- 
alogue of inammals in the American Mu- 
seum of Natural History (Lawrence, 
1993). For systematic consistency, the tax- 



Type Specimens of Recent Mammals • Helgen and McFadden 97 



onomic judgments at the species-level by 
the authors of the chapters in Mammal 
Species of the World (Wilson and Reeder, 
1993) are largely adhered to. However, in 
many cases alternate views are explored, 
and subsequent work by other authors is 
noted. The sequence of mammalian orders 
presented here is as follows: Didelphimor- 
phia, Dasyuromorphia, Peramelia, Cingu- 
lata, Afrosoricida, Rodentia, Lagomoi-pha, 
Scandentia, Primates, Lipotyphla, Chirop- 
tera, Artiodactyla, Carnivora, and Cimoles- 
ta. This sequence represents the ongoing 
understanding of higher mainmalian rela- 
tionships being produced by research in 
molecular phylogenetics (Waddell et al., 
1999) as well as paleontological studies 
(McKenna and Bell, 1997). A number of 
these names are rather nontraditional; use 
of the names Cingulata (for armadillos) 
and Cimolesta (for pangolins) at ordinal 
rank follows McKenna and Bell (1997). Af- 
rosoricida is used for an order including 
tenrecs and golden-moles (following Stan- 
hope et al, 1998: 9971-9972). We use the 
ordinal name Lipotyphla in a restricted 
sense to refer to the Recent families Ne- 
sophontidae, Solenodontidae, Soricidae, 
and Talpidae (others have used the term 
"Eulipotyphla," e.g., Waddell et al, 1999). 
Cetaceans are included here within the or- 
der Artiodactyla (an assemblage often re- 
ferred to as "Cetartiodactyla" in recent lit- 
erature; see Graur et al., 1997). 

To avoid unnecessaiy complexity, or- 
ders, families, and genera are the only 
ranks above the level of species that are 
listed, except for the large family Muridae, 
for which subfamilial distinctions are pro- 
vided, in alphabetical order. Within each 
order, the sequence of families generally 
follows Simpson (1945), but Wilson and 
Reeder's (1993) order of rodent families 
and Simmons' (1998: 12) arrangement of 
the bats are observed. Within each genus, 
taxa are presented in alphabetical order by 
original name. 

The format of this catalogue is as fol- 
lows, with all the following information 
provided when possible: 



Original binomen. Name of describer, date 

of description. 

Citation of original publication. 

= Presently used name, if different from 

original. Citation of publication in which 

this name combination was initially used 

for this taxon. 

Type Category. Number of specimen.^ 
Preparation of specimen (skin, skull, alco- 
hol, etc.), age and sex. 

Locality. Type locality. Date of collec- 
tion. 

Collector. Name of collector. Original 
number of specimen. 

Condition. Current condition of the 
type material. 

Type Series. Any paratypes, paralecto- 
types, or additional syntypes in existence 
are mentioned, with their preparation, sex, 
and age. 

Comments. Additional comments re- 
garding the systematic status or the histoiy 
of the specimen. 

TAXA INCLUDED IN THIS CATALOGUE 

Unlike Allen's original type catalogue, fos- 
sil mammals are not considered here, but 
type specimens of Recent mammals known 
only from subfossil remains are discussed. 

Several type specimens of Recent mam- 
mals that were included in Allen's cata- 
logue are not considered here as name- 
bearing types. A number of syntype series 
in the MCZ have since Allen's time been 
rendered paralectotypes by the designa- 
tion of a lectotype preserved in another 
institution; these are discussed in the sec- 
tion below on paralectotypes. Additionally, 
MCZ 14929, listed as a "cotype" of Nyc- 
teris revoili Robin, 1881 by G. M. Allen 
(1931: 235), is a paratype rather than a 
syntype and is not considered here. 



- Mammal specimens in the MCZ bear any of three 
kinds of numbers. A number preceded by "MCZ" can 
be found in the general collection in the Mammal 
Department. A number preceded by a "B" is part of 
the collection of E. A. and O. Bangs, also housed in 
the Mammal Department. A number preceded by 
"VP" designates that the specimen is stored in the 
Vertebrate Paleontology Department. 



98 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



One other enigmatic specimen in the col- 
lection deseives some mention. When it was 
received by die Mammal Department, Guil- 
laume Grandidiers personal collection con- 
tained specimens marked as types for 16 
taxa, described by either Grandidier or M. 
L. Lavauden. Fifteen of these holotypes are 
discussed witliin this catalogue in separate 
accounts. The remaining specimen, a fioiit 
bat, MCZ 45073, is marked exactly like the 
odier type specimens, and both its sldn and 
its skull tags bear the name "Eidolon saka- 
lava nov. spec, G. Grandidier" in die script- 
ed handwriting typical of Grandidiers spec- 
imens. The locality given for the specimen 
is Ankavandra, in west-central Madagascar. 
Though this specimen has been curated as 
a type specimen, I (KMH) can find no pub- 
lished description of diis taxon or any ref- 
erence to diis name in the literature. This 
name is dierefore invalid, barring a discov- 
eiy in the future diat it was indeed pub- 
lished and has since been overlooked. What- 
ever die published status of the name, it ap- 
pears to me diat diis specimen should be 
considered as a young specimen of Eidolon 
dupreanum rather dian a distinct species, 
and discussion of this mysterious binomial 
here should in no way be constiaied as a 
formal description of a new taxon. 

ABBREVIATIONS 

Abbreviations are used in the text to 
designate the following institutions. 

AMNH American Museum of Natural 
Histoiy, New York 

BMNH Natural Histoiy Museum, Lon- 
don 

BSNH Boston Society of Natural His- 
tory, Boston 

FMNH Field Museum of Natural His- 
tory, Chicago 

MNHN Museum National d'Histoire 
Naturelle, Paris 

MVZ Museum of Vertebrate Zoology, 

Berkeley, California 

RMNH Rijksmuseum van Natuurlijke 
Historic, Leiden 



USNM 



YPM 



National Museum of Natural 
Histoiy Washington, D.C. 
Yale Peabody Museum of Nat- 
ural History, New Haven, Con- 
necticut 



ACKNOWLEDGMENTS 

We would like to thank the faculty and 
staff of the Mammal Department at the 
MCZ — A. W. Crompton, Andrew Biewe- 
iier, Judith Chupasko, Jane Harrison, and 
especially Maria Rutzmoser for access to 
the type specimens. Maiy Sears, Timothy 
McNeece, and Ronnie Broadfoot at the 
Ernst Mayr Libraiy, HaiA^ard University, 
were extremely helpful in pinning down 
difficult references. We also thank Alison 
Pine for allowing us to make use of the 
MCZ Ornithology Department locality 
references; Carolyn Kirdahy of the Boston 
Museum of Science, w^lio granted us ac- 
cess to the archives of the Boston Society 
of Natural History; and the many individ- 
uals associated with other museums who 
assisted by answering any questions that 
arose. Finally, we are thankful for the sug- 
gestions we received from two anonymous 
reviewers, which helped us improve this 
catalogue. 

PARALECTOTYPE SERIES 

Hesperomys eremicus Baird, 1858 =Peromyscus er- 
emicus eremicus (Baird, 1858). Lectotype, USNM 
2575, designated by Osgood (1909: 241). MCZ 
4310 and 5273 are paralectotypes. 

Neotoma fuscipes Baird, 1858 =Neotoinafuscipesfus- 
cipes Baird, 1858. Lectotype, USNM 22026, des- 
ignated by Lyon and Osgood (1909: 99). MCZ 4336 
and 5264 are paralectotypes. 

Mus bairdii Hoy and Kennicott, 1857 =Peromyscus 
maniculatus bairdii (Hoy and Kennicott, 1857). 
Lectotype, number 750 in the Collection of the 
Academy of Natural Sciences of Philadelphia, des- 
ignated by Osgood (1909: 80). MCZ 8073 is a par- 
alectotype. 

Pteropus lanigera [sic] H. Allen, 1890 =Pteropus in- 
stdaris Hombron and Jacquinot, 1842. The cor- 
rected spelling of the original name is Pteropus lan- 
iger (see Andersen 1912: 297). H. Allen based the 
description of Pteropus laniger on two syntypes, 
USNM 19066 (skin)/37815 (skull) and MCZ 7023, 
a skin. Andersen (1912: 297-298) was unaware of 



Type Specimens of Recent Mammals • Helaen and McFadden 



99 



the whereabouts of this latter specimen and based 
his evaluation of the systematic status of P. laniger 
solely on the USNM specimen in his taxonomic 
review of the Megachiroptera. Because it is rep- 
resented by both a skin and a skull in good con- 
dition and because it has been used in past taxo- 
nomic treatments, USNM 19066/37815 is hereby 
designated as the lectotype of Pteropus laniger to 
ensure consistency between past and future taxo- 
nomic treatments of this name. MCZ 7023 is thus 
a paralectotype. The type locality of laniger is the 
Caroline Islands, as emended by Andersen (1912: 
298), not Samoa, as originally described. 

Sciunis castanotus Baird, 1855 and Sciurus castan- 
onotus Baird, 1858 =Scmnis aherti aherti Wood- 
house, 1853. Baird (1858: 266) noted that the name 
castanotus, used in his original description of this 
taxon, was a misprint for castanonotus. Lectotype, 
USNM 121/1107, designated by Lyon and Osgood 
(1909: 183). Though not explicitly stated by Lyon 
and Osgood, this specimen should serve as a lec- 
totype for both names (Sciunis castanotus Baird, 
1855 and Sciunis castanonotus Baird, 1858), for 
ttixonomic consistency. MCZ 4692 is a paralecto- 
type. 

Spennophiliis obsoletus Kennicott, 1863 =Spenno- 
philus spilosoma obsoletus Kennicott, 1863. Lec- 
totyjoe, USNM 3222/27998, designated by A. H. 
Howell (1938: 130). See G. M, Allen (1931: 252) 
for a list of paralectotypes (then considered synty- 
pes) in the MCZ. 

Spennophiliis parnji van kodiacensis J. A. Allen, 1874 
= Spennophiliis parnji kodiacensis J. A. Allen, 
1874. Lectotype, USNM 9242/38543, designated 
by A. H. Howell (1938: 103). See G. M. Allen 
(1931: 252) for a list of paralectotypes (then con- 
sidered syntypes) in the MCZ. 

Spennophiliis tridecemlineatus van pallidus ]. A. Al- 
len, 1874 = Spennophiliis tridecemlineatus pallidus 
J. A. Allen, 1874. Lectotype, USNM 16237, des- 
ignated by A. H. Howell (1938: 112). See G. M. 
Allen (1931: 253) for a Ust of paralectotypes (then 
considered syntypes) in the MCZ. 

Tamias quadrivittatus var. pallidus J. A. Allen, 1874 
= Tamias minimus pallidus J. A. Allen, 1874. Lec- 
totype, USNM 11656/38311, designated by Gary 
(1906: 88). G. M. Allen (1931: 255) provided a hst 
of paralectotypes. 

ACCOUNTS OF NAME-BEARING TYPE 
SPECIMENS 

Order DIDELPHIMORPHIA Gill, 1872 

Family CALUROMYIDAE Kirsch and Reig, 
1977 

Genus CALUROMYSJ. A. Allen, 1900 

Philander cicur Bangs, 1 898k 



Proc. Biol. Soc. Washington, 12: 161, 10 

August. 

= Caluromys lanatus cicur {Bangs, 1898). 

See Cabrera (1958: 2). 

Holotype. B8114. Skin and skull. Adult female. 

Locality. Colombia: (Magdalena), Santa Marta 
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 27 
March 1898. 

Collector. W. W. Brown, Jr. Original number 123. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Tijpe Series. 3 paratypes; B8036, skin and skull, 
adult male; B8115, skin and skull, adult male; 
B8116, skin and skull, adult male. 

Family MARMOSIDAE Hershkovitz, 1992 

Genus MARMOSA Gray, 1821 

Marmosa robinsoni Bangs, 18981 

Proc. Biol. Soc. Washington, 12: 95, 30 

April. 

= Marmosa robinsoni robinsoni Bangs, 

1898. See Cabrera (1958: 24). 

Holotype. B7749. Skin and skull. Adult male. 

Locality. Venezuela: (Nueva Esparta), Margarita 
Island. 12 July 1895. 

Collector. W. Robinson. Original number 506. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 2 paratypes, both in the USNM; 
USNM 63209, skin and skull, adult male; USNM 
63210, skin and skull, adult female. 
Comments. M. robinsoni was considered a valid 
species by Gardner (1993: 18) and Nowak (1999: 
21). 

IVIarmosa mitis Bangs, 1 898k 

Proc. Biol. Soc. Washington, 12: 162, 10 

August. 

= Marmosa robinsoni robinsoni Bangs, 

1898. See Cabrera (1958: 24). 

Holotype. B8123. Skin and skull. Adult male. 

Locality. Colombia: (Magdalena), Santa Marta 
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 25 
March 1898. 

Collector. W. W. Brown, Jr. Original number 91. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Tijpe Series. 26 paratypes; B8117-B8122, B8124- 
B8143; all represented by skin and skull, 15 fe- 
males and 11 males. 5 paratypes are no longer in 
the MCZ (B8118 is at Wellesley College, B8124 
and B8141 are at FMNH, and B8136 and B8138 
are at USNM). 



100 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Marmosa fulviventer Bangs, 1901b 
Amer. Nat., 35: 632, 22 August. 
= Marmosa robinsoni fulviventer Bangs, 
1901. See Handley (1966: 775). 

Holotype. B8435. Sldn and skull. Adult male. 

Locality. Panama: (Panama), Gulf of Panama, San 
Miguel Island. 28 April 1900. 

Collector. W. W. Brown, Jr. Original number 123. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; B8436, skin and skull, fe- 
male (exchanged to FMNH in 1931); B8437, skin 
and skull, female; B8438, skin and skull, male. 
Comments. M. r. fulviventer was retained as a valid 
subspecies by Hall (1981: 14) and O'Connell (1983: 
1). 

Family DIDELPHIDAE Gray, 1821 

Genus D/DELPH/S Linnaeus, 1758 

Didelphis marsupialis particeps Goldman, 

1917 

Proc. Biol. Soc. Washington, 30: 107, 23 

May. 

Holotype. B8439. Skin and skull. Adult male. 

Locality. Panama: (Panama), Gulf of Panama, San 

Miguel Island. 8 May 1900. 

Collector. W. W. Brown, Jr. Original number 165. 

Condition. Skin and skull complete. 

Type Series. 1 paratype; B8440, skin and skull, 

adult female. 

Comments. Retained as a valid subspecies by Hall 

(1981: 4). 

Didelphis virginiana pigra Bangs, 1 898b 
Proc. Boston Soc. Nat. Hist., 28: 172, 15 
March. 

Holotype. B3500. Sldn and skull. Adult female. 
Locality. (United States): Florida, Brevard Gounty, 
Oak Lodge, East Peninsula opposite Micco. 31 Jan- 
uaiy 1895. 
Collector. O. Bangs. 
Condition. Skin and skull complete. 
Type Series. 11 paratypes; all represented by skin 
and skull, most still in the MGZ. 
Comments. Retained as a valid subspecies by Hall 
(1981: 5) and McManus (1974: 1). The type de- 
scription lists 31 January 1896 as the date of col- 
lection, but the date is written as "January 31, 
1895" on the original specimen label and in Bangs" 
accession catalogue. 

Order DASYUROIVIORPHIA Gill, 1872 
Family DASYURIDAE Goldfuss, 1820 
Genus ANTECHINUS Macleay, 1841 



Antecfiinus may eh misim Tate, 1947 

Bull. Amer. Mus. Nat. Hist., 88: 130, 20 

February. 

= Antecfiinus naso misim Tate, 1947. See 

Laurie and Hill (1954: 7). 

Holotype MGZ 29924. Skin and skull. Adult male. 
Locality. Papua New Guinea: Morobe Province, 
Mount Misim ( = Missim), 5,850 ft (1,784 m). 14 
April 1933. 
Collector H. Stevens. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; MGZ 29923, sldn and 
skull, adult female. 

Comments. Tate, in the original description, erro- 
neously listed 24 April 1933 as the date of collec- 
tion. A. n. misim was retained as a valid subspecies 
by Flanneiy (1995a: 80), who also noted that the 
New Guinean species assigned to the genus Ante- 
chinus are not closely related to the Australian spe- 
cies of that genus and will be reassigned at the 
generic level pending a full taxonomic revision of 
the group. 

Genus IVIYOICTIS Gray, 1858 

Myoictis melas wavicus Tate, 1 947 

Bull. Amer. Mus. Nat. Hist., 88: 140, 20 

February. 

= Myoictis melas wallacei Gray, 1858. See 

Flanneiy (1990: 56). 

Holotijpe. MGZ 28082. Skin and skull. Adult male. 
Locality. Papua New Guinea (S. Morobe), Wau, 
3,800 ft (1,159 m). 27 March 1932. 
Collector H. Stevens. Original number 1. 
Condition. Sldn and skull complete. 
Type Series. Holot}'pe only. 

Comments. Stevens' original field label bears the 
following lament — "To my intense anguish I failed 
to retrieve the female, shot on recumbent, decayed 
log in undergrowth." 

Order PERAMELIA Ameghino, 1889 

Family PERORYCTIDAE Groves and 
Flannery, 1990 

Genus ECHYMIPERA Lesson, 1842 

Suillomeles hispida G. M. Allen and 

Barbour, 1909 

Proc. New England Zool. Club, 4: 44, 12 

July. 

= Echymipera kalubu kalubu (Fischer, 

1829). See Laurie and Hill (1954: 11). 

Holotype. MGZ 7006. Skin and skull. Adult. 



Type Specimens of Recent Mammals • Helaen and McFadden 



101 



Localitij. (Indonesia): Dutch New Guinea ( = Ii-ian 

Jaya), Doreh Bay, Manokwari, "not far from the 

foot of Mt. Arfak." 23 Februaiy 1907. 

Collector. T. Barbour. 

Condition. Skin complete. Skull partial (occipital 

region and two posterior upper molars on each side 

missing). 

Type Serie.s. Holotype only. 

Comments. S. hispida is the type species of Suil- 

lomeles G. M. Allen and Barbour, 1909. 



Echymipera rufescens austral is Tate, 1948 
Bull. Amer. Mus. Nat. Hist., 92: 334, 25 
November. 



Holotype. MCZ 29214. Skin and skull. Adult male. 
Locality. Australia: Queensland, Cape York, near 
Coen, east slope of Mcllwraith Ranges. Rocky Riv- 
er, "Rocky Scrub." 20 June 1932. 
Collector. R J. Darlington, Jr., Hanard Australian 
Expedition. Original number 209. 
Condition. Skin complete, but tip of tail worn. 
Skull complete. 
Type Series. Holotype only. 

Comments. Tate (1952: 582) reported that the ex- 
act type locality is "in the dense rain forests of the 
upper Nesbit River on the east slopes of the 
Mcllwraith Range." Retained as a valid subspecies 
by Flanneiy (1995a: 111). Darlington noted in his 
field notebook, regarding this specimen, "Animal 
about the fattest I have skinned — I shall dream of 
it!" 



Order CINGULATA llliger, 1911 

Family DASYPODIDAE Gray, 1821 

Genus D/ASVPL/S Linnaeus, 1758 

Dasypus novemcinctus hoplites G. M. 

Allen, 1911a 

Bull. Mus. Comp. Zool., 54: 195, July. 



Holotype. MCZ 8116. Skin, skull, and postcranial 
skeleton. Adult female. 

Locality. Grenada: hills back of Gouyave. 7 Sep- 
tember 1910. 

Collector G. M. Allen. Original number 26. 
Condition. Skin, skull, and postcranial skeleton 
complete. 

Type Series. 2 paratypes; MCZ 8117, skin and skull, 
adult male; MCZ 8118, skin and skull, adult male. 
Comments. Retained as a valid subspecies by Hall 
(1981: 283) and McBee and Baker (1982: 1). 



Order AFROSORICIDA Stanhope et a!., 
1998 

Family CHRYSOCHLORIDAE Gray, 1825 

Genus CHRYSOCHLORIS Lacepede, 
1799 

Chlorotalpa tropicalis G. M. Allen and 

Loveridge, 1927 

Proc. Boston Soc. Nat. Hist., 38: 418, 23 

December. 

= Chrysochloris stuhlmanni tropicalis (G. 

M. Allen and Loveridge, 1927). See 

Meester (1974: 3). 

Holotype. MCZ 22435. Skin and skull. Adult female. 
Locality. Tanganyika Territory (=Tanzania): Ulu- 
gui"u Mountains, Bagilo. 5 October 1926. 
Collector A. Loveridge. 

Condition. Skin complete. Skull slightly damaged 
(coronoid processes missing from both mandibular 
rami). Mandible disarticulated. 
Type Series. Holotyj^De only. 

Comments. Included in Chrysochloris stuhlmanni 
by Meester (1974: 3) but recognized as distinct by 
Simonetta (1968: 42). Hutterer (1993: 75) noted 
that the systematic status of tropicalis merits fur- 
ther study. 

Family TENRECIDAE Gray, 1821 

Genus GEOGALE Milne-Edwards and A. 
Grandidier, 1872 

Cryptogale australis G. Grandidier, 1928 

Bull. Mus. Hist. Nat. Paris, 34: 64, 26 

Januaiy. 

= Geogale aurita Milne-Edwards and A. 

Grandidier, 1872. See Genest and Petter 

(1975: 3). 

Holotype. MCZ 45057. Skull fragments. 

Locality. Madagascar: (Toliary), south of Fort Dau- 
phin (=Tolanaro), Andrahomana grotto. 1927. 
Collector R. Decary. 

Condition. MCZ 45047 includes 18 partial crania 
and 8 mandibular rami. Fragmentary. 
Type Series. All the type material of C australis 
bears a single accession number. 
Comments. C. australis is the type species of the 
genus Cn/pto^ale G. Grandidier, 1928. 

Geogale aurita orientalis G. Grandidier 

and Petit, 1930 

Faune des Colonies Frangaises, 4: 446. 

Holotype. MCZ 45660. Body in alcohol, cranium sep- 
arate. 



102 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Locality. Madagascar: (Toamasina), east coast, Fe- 
iierive ( = Fenoarivo Atsinanana). April 1928. 
Collector. R. Decaiy. Original number 12. 
Condition. Alcoholic, cranium complete. 
Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Ge- 
nest and Fetter (1975: 3). The holotype of G. a. 
orientalis seems to be the only record of Geogale 
aurita from the east coast of Madagascar. 

Genus /W/CROG/\/.E Thomas, 1882 

Microgale decaryi G. Grandidier, 1928 

Bull. Mus. Hist. Nat. Paris, 34: 69, 26 

January. 

= Microgale principula Thomas, 1926. See 

MacPhee (1987a: 9). 

Holotype. MCZ 45049. Cranium. Adult. 

Locality. Madagascar: (Toliary), south of Fort Dau- 
phin (=Tolanaro), Andrahomana caves. 1926. 
Collector. R. Decary. 

Condition. Cranium partial (back of cranium miss- 
ing posterior to parietals). 

Type Series. Paratype material consists of MCZ 
45408, which includes 3 partial crania and 5 man- 
dibular rami, and MAD- 1649, a partial skull in the 
collections of the Institut de Paleontologie, 
MNHN. 

Microgale drouhardi G. Grandidier, 1934 
Bull. Mus. Hist. Nat. Paris, 6: 474, 29 
November. 

Holotype. MCZ 45034. Body in alcohol, skull extract- 
ed. Juvenile female. 

Locality. Madagascar: (Antsiranana), east coast, Di- 
ego-Suarez (= Antsiranana). May 1934. 
Collector M. E. Drouhard. Original number A. 
Condition. Alcoholic, skull complete. 
Type Series. 6 paratypes; MCZ 46007-MCZ 46012, 
all in alcohol. MCZ 46017, represented by a skull 
and postcranial skeleton, was collected at the same 
time and place as the type series but is not men- 
tioned in the original description. 
Comments. MCZ 45034 represents an immature 
animal (MacPhee 1987a: 7), not an adult as claimed 
in the original description. MacPhee (1987a: 9) 
synonyinized drouhardi with Microgale cowani, an 
approach followed by Hutterer (1993: 71), but Jen- 
kins et al. (1997: 6) argued that M. droidiardi is a 
distinct species. 

Microgale parvula G. Grandidier, 1934 
Bull. Mus. Hist. Nat. Paris, 6: 476, 29 
November. 

Holotype. MCZ 45465. Body in alcohol, skull extract- 
ed. Juvenile male. 



Locality. Madagascar: (Antsiranana), east coast, Di- 
ego-Suarez (= Antsiranana). May 1934. 
Collector. M. Drouhard. 
Condition. Alcoholic, skull complete. 
Tijpe Series. Holotype only. 

Comments. MCZ 45465 represents an immature 
animal (MacPhee 1987a: 7), not an adult as claimed 
in the original description. The holotype of Micro- 
gale pulla Jenkins, 1988 actually represents an 
adult specimen of M. parvula (Jenkins et al., 1996: 
204). Considered a valid species by Hutterer (1993: 
71) and Nowak (1999: 190). 



Microgale prollxacaudata G. Grandidier, 

1937 

Bull. Mus. Hist. Nat. Paris, 9: 348, 25 

November. 

= Microgale longicaudata Thomas, 1882. 

See MacPhee (1987a: 9). 



Holotype. MCZ 45035. Body in alcohol, skull extract- 
ed. Juvenile. 

Locality. Madagascar: (Antsiranana), east coast, Di- 
ego-Suarez (= Antsiranana). May 1934. 
Collector M. Drouhard. 

Condition. Alcoholic; skull partial (left tympanic 
bulla missing). Mandible disarticulated. 
Type Series. 1 paratype; MCZ 46020, in alcohol. 
Comments. This specimen represents an immature 
animal (MacPhee 1987a: 8), not an adult as claimed 
in the original description. 



Paramlcrogale occldentalls G. Grandidier 
and Petit, 1931 

Bull. Soc. Zool. France, 56: 129, 15 June. 
= Microgale brevlcaudata G. Grandidier, 
1899. See MacPhee (1987a: 9). 



Holotype. MCZ 45047. Body in alcohol, skull extract- 
ed. Juvenile male. 

Locality. Madagascar: (Antananarivo), northwest of 
Maintirano, Andriafeuelo. 1930. 
Collector M. A. de la Rue. 
Condition. Alcoholic, skull complete. 
Type Series. Holotype only, but see comments. 
Comments. P. occidentalis is the type species of the 
genus Paramicrogale G. Grandidier and Petit, 
1931. This specimen represents an immature ani- 
mal (MacPhee 1987a: 7), not an adult as claimed 
in the original description. The original description 
mentions only a single specimen but describes ex- 
tensively the skeleton of P. occidentalis. This is puz- 
zling, as the skeleton has not been extracted from 
MCZ 45047 (MacPhee 1987a: 7). 



Type Specimens of Recent Mammals • Helgen and McFadden 103 



Genus SETIFER Fronep, 1806 

Dasogale fontoynonti G. Grandidier, 

1930a 

Bull. Acad. Malgache, n. sen, 11: 85 (for 

1928). 

= Setifer setosus {Schreber, 1777). See 

Poduschka and Poduschka (1982: 261). 



Holotype. MCZ 45016. Skin, skull, and postcranial 
skeleton. Juvenile. 

Localitij. Madagascar: east coast. 1917. 
Collector. Received by G. Grandidier from the 
Academic Malgache in 1917. 

Condition. Sldn represented by a small patch of 
lur. Skull partial (occipital region missing). Postcra- 
nial skeleton complete, partially articulated. 
Tijpe Series. 1 paratype; MCZ 45532, in alcohol. 
Comments. Walker (1975: 110) commented that 
"the only specimen known of D. fontoynonti ... is 
in the Paris Museum"; actually, the only material 
attributed to Dasogale is in the MCZ, a fact first 
noted in publication by Poduschka and Poduschka 
(1982: 253). Dasogale was often considered to be 
an extremely rare or recently extinct species until 
Poduschka and Poduschka (p. 261) and MacPhee 
(19S7b: 135) demonstrated that the holotype is 
probably a juvenile Setifer .setosus. D. fontoynonti 
is the type species of the genus Dasogale G. Gran- 
didier, 1930. 



Order RODENTIA Bowdich, 1821 

Family APLODONTIDAE Brandt, 1855 

Genus APLODONTIA Richardson, 1829 

Aplodontia californica columbiana Taylor, 

1916 

Univ. California Publ. Zool., 12: 499, 6 

May. 

= Aplodontia rufa rainieri Merriam, 1899. 

See Dalquest (1948: 369). 



Holotype. B1899. Sldn and skull. Adult male. 

Locality. (Canada): British Columbia, Hope, 

Roabs Ranch. 14 June 1894. 

Collector W. C. Colt. Original number 479. 

Condition. Skin and skull complete. 

Type Series. 8 paratypes; B1892-B1898, B1900; all 

represented by skin and skull, 3 females and 5 

males. 



Family SCIURIDAE Fischer de Waldheim, 
1817 

Genus CALLOSCIURUS Gray, 1867 

Callosciurus baluensis medial is G. M. 

Allen and Coolidge, 1940 

Bull. Mus. Comp. Zool., 87: 156, 31 

December. 

Holotype. MCZ 22265. Skin and skull. Adult female. 
Locality. (Indonesia): Dutch Borneo, (Kalimantan), 
Mount Tibang (possibly =Bukit Tungun). 1925. 
Collector. E. Mjoberg. Original number 6. 
Condition. Skin complete. Skull partial (right pa- 
rietal, right tympanic bulla, and lachiymal bro- 
ken — bulla present). Mandible disarticulated. 
Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Med- 
way (1977: 90). 

Callosciurus ferruginous primus G. M. 

Allen and Coolidge, 1940 

Bull. Mus. Comp. Zool, 87: 157, 31 

December. 

= Callosciurus erythraeus (Pallas, 1778). 

See Corbet and Hill (1992: 283). 

Holotype. MCZ 35352. Skin and skull. Adult female. 
Locality. Siam (=Thailand): Mae Wan River near 
Doi, Mount Souket ( = Saket), 1,500 ft (458 m). 20 
Febiaiary 1937. 

Collector J. A. Griswold, Jr., Asiatic Primate Ex- 
pedition. Original number 5. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; MCZ 35353, skin and 
skull, adult male. 

Sciurus castaneoventris haemobaphes G. 

M. Allen, 1912c 

Proc. Biol. Soc. Washington, 25: 177, 24 

December. 

= Callosciurus erythraeus haemobaphes 

(G. M. Allen, 1912). See Hayman and 

Holt (1940: 359). 

Holotype. MCZ 13693. Skin and skull. Male. 

Locality. China: southeastern Yunnan, Chih-ping 
( = Shiping). 26 Februaiy 1911. 
Collector Kobayashi Collection. Original number 
46. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Only the holotype is mentioned in the 
original publication, but Allen had four other spec- 
imens of these squirrels at the time of description 
(MCZ 13692, 13694-13696; all represented by skin 
and skull, 3 males, 1 female). 



104 Bulletin Museum of Comparative Zoolog^y, Vol. 157, No. 2 



Genus DREMOMYS Heude, 1 898 

Dremomys pernyi flavior G. M. Allen, 

1912c 

Proc. Biol. Soc. Washington, 25: 178, 24 

December. 

Holottjpe. MCZ 13691. Skin and skull. Male. 

Locality. China: southeastern Yunnan, Mongtz 
( = Mengzi). 1911. 

Collector. Kobayashi Collection. Original number 
6/8. 

Condition. Skin complete. Skull partial (squamo- 
sals missing, supraoccipital damaged, right jugal 
missing, palatine missing, tympanic bulla dam- 
aged). 
Type Series. Holoty]^)e only. 

Dremomys senex G. M. Allen, 1912b 

Mem. Mus. Comp. ZooL, 40: 229, 

August. 

= Dremomys pernyi senex G. M. Allen, 

1912. See Hayman and Holt (1940: 382). 

Holotype. MCZ 7582. Skin and skull. Adult female. 
Locality. China: Hupeh ( = Hubei), Ichanghsien, 
Nantou. 5 Februaiy 1909. 

Collector W. R. Zappey. Original number .373. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paraty^je; MCZ 7583, skin and skull, 
adult male. 

Genus FUNISCIURUS Trouessart, 1880 

Funisciurus pyrrhopus victoriae G. M. 

Allen and Loveridge, 1942 

Bull. Mus. Comp. Zool., 8: 180, 

February. 

= Funisciurus pyrriiopus al<l<a De Winton, 

1899. See Amtmann (1975: 8). 

Holotype. MCZ 39199. Skin and skull. Adult male. 
Locality. Uganda: Toro, Kibale Forest, 4,200 ft 
(1,281 m). 16 December 1938. 
Collector A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Genus GLAUCOIVIYS Thomas, 1908 

Sciuropterus alpinus bangsi Rhoads, 1897 
Proc. Acad. Nat. Sci. Philadelphia, 1897, 
p. 321 (footnote), July. 
^Glaucomys sabrinus bangsi (Rhoads, 
1897). See A. H. Howell (1918: 38). 

Holotype. B6959. Sldn and skull. Adult male. 

Locality. (United States): Idaho, Idalio County. 
March 8, 1897. 



Collector Harbison and Bargamin. 
Condition. Skin and skull complete. 
Type Series. 1 paratype; B6960, skin and skull, 
adult male. 

Comments. G. s. bangsi was retained as a valid sub- 
species by Hall (1981: 450). 

Sciuropterus alpinus lascivus Bangs, 

1899J 

Proc. New England Zool. Club, 1: 69, 31 

July. 

= Glaucomys sabrinus lascivus (Bangs, 

1899). See A. H. Howell (1918: 55). 

Holotype. B9186. Skin and skull. Adult female. 
Locality. (United States): California, El Dorado 
County, Tallac. 28 August 1898. 
Collector W. VV. Price and P. O. Simons. Original 
number 1722. 

Condition. Sldn and skull complete. 
Type Series. 2 paratypes, B9187, skin and skull, fe- 
male; B9188, sldn and skrdl, female. 
Comments. G. s. la.scivns was retained as a valid 
subspecies by Hall (1981: 451). 

Sciuropterus sabrinus makkovikensis 

Sornborger, 1900 

Ottawa Nat., 14: 48, 6 June. 

= Glaucomys sabrinus makkovikensis 

(Sornborger, 1900). See A. H. Howell 

(1918: 34). 

Syntypes. MCZ 10476: Skin and skull. Adult. MCZ 
10477: Skin and skull. Adult. MCZ 10478: Skin and 
skull. Adult. 

Locality. (Canada): Labrador Peninsula, Makkovik. 
1899. 

Collector. W. W. Perrett. Original number 1540. 
Condition. MCZ 10476: Skin complete. Skull par- 
tial (base of skull missing). Mandible disarticulated. 
MCZ 10477: Skin partial (tail broken but present). 
Skull partial (base of skull missing). Mandible dis- 
articulated. MCZ 10478; Skin partial (missing left 
hind foot and tail). Skull partial (occipital chipped). 
Mandible disarticulated. 
Type Series. 3 syntypes only. 

Comments. Sornborger's original description was 
based on three specimens, original numbers 1540, 
1541, and 1542, now MCZ 10476, 10477, 10478, 
respectively. G. .$. makkovikensis was retained as a 
valid subspecies by Hall (1981: 453). 

Sciuropterus silus Bangs, 1896J 

Proc. Biol. Soc. Washington, 10: 163, 28 

December. 

= Glaucomys volans volans (Linnaeus, 

1758). See A. H. Howell (1918: 20). 

Holotype. B4931. Sldn and skull. Adult male. 

Locality. (United States): West Virginia, Greenbri- 



Type Specimens of Recent Mammals • Helpen and McFodden 



105 



er County, White Sulphur Springs, top of Katis 
Mtn. 3,200 ft (976 m). 2 September 1895. 
Collector. T. Surber. Original number 19. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Sciuropterus volans querceti Bangs, 1 896j 

Proc. Biol. Soc. Washington, 10: 166, 28 

December. 

= Glaucomys volans querceti (Bangs, 

1896). See A. H. Howell (1918: 26). 

Holotype. B2451. Skin and skull. Adult female. 
Locality. (United States): Florida, Citrus County, 
Citronelle. 17 September 1894. 
Collector F. L. Small. Original number 1363. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 2 paratypes; B2452, skin and skull, 
adult male; B2453, skin and skull, adult female. 
Comments. G. v. querceti was retained as a valid 
subspecies by Hall (1981: 449). 

Genus HYLOPETES Thomas, 1 908 

Pteromys (Hylopetes) alboniger orinus G. 

M. Allen, 1940 

The Mammals of China and Mongolia, 

Natural Histoiy of Central Asia, 11: 723, 

3 September. 

= Hylopetes alboniger alboniger (Hodgson, 

1836). See Ellerman and Morrison-Scott 

(1951: 469). 

Holotype. MCZ 28086. Skin and skull. Adult female. 
Locality. China: Yunnan, Likiang Range ( = Li- 
jiang), 7,800 ft (2,379 m). December 1931. 
Collector J. F Rock. 

Condition. Skin complete. Skull partial (most of 
right tympantic bulla missing, supraoccipital dam- 
aged). Mandible disarticulated. 
Type Series. Allen examined 10 specimens in ad- 
dition to the type, including 4 specimens from the 
BMNH, 3 from AMNH, and MCZ 28087, skin and 
skull, an unsexed adult. 

Pteromys phayrei anchises G. M. Allen 

and Coolldge, 1940 

Bull. Mus. Comp. Zool., 87: 153, 31 

December. 

= Hylopetes phayrei anchises (G. M. Allen 

and Coolldge, 1940). See Ellerman and 

Morrison-Scott (1951: 469). 

Holotype. MCZ 35776. Skin and skull. Adult male. 
Locality. Siam (=Thailand): (Chiang Mai) Mount 



Angka ( = Doi Inthanon), 4,300 ft (1,312 m). 27 
February 1937. 

Collector. J. A. Griswold, Jr., Asiatic Primate Ex- 
pedition. Original number 24. 

Condition. Skin and skvdl complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; MCZ 35775, skin and 
sk-ull, adult female; MCZ 35777, skin and skull, 
adult male; MCZ 35778, skin and skull, subadult 
male. 

Genus MARMOTA Blumenbach, 1779 

Arctomys flaviventer avarus Bangs, 1899] 

Proc. New England Zool. Club, 1: 68, 31 

July. 

= Marmota flaviventris avara (Bangs, 

1899). See A. H. Howell (1915: 41). 

Holotype. B7299. Skin and skull. Juvenile female. 
Locality. (Canada): British Columbia, Okanagan. 
17 July 1897. 

Collector A. C. Brooks. Original number 969. 
Condition. Skin and skadl complete. 
Type Series. 2 paratypes; B7298, skin and skull, ju- 
venile male; B7300, skin and skull, juvenile female. 
Comments. M. f. avara was retained as a valid sub- 
species by Hall (1981: 371). 

Arctomys ignavus Bangs, 1899d 

Proc. New England Zool. Club, 1: 13, 28 

February. 

= Marmota monax ignava (Bangs, 1899). 

See A. H. Howell (1915: 29). 

Holotype. B7971. Skin and skull. Adult male. 

Locality. (Canada): Labrador Peninsula, Black Bay. 
13 July 1898. 
Collector. E. Doane. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 5 paratypes; B7968-B7970, B7972, 
B7973 (juvenile); all represented by skin and skull, 
4 females and 1 male. 

Comments. M. m. ignava was retained as a valid 
subspecies by Hall (1981: 370) and Kwiecinski 
(1998: 1). 

Genus MICROSCIURUS J. A. Allen, 1895 

Sciurus (Microsciurus) browni Bangs, 

1902b 

Bull. Mus. Comp. Zool., 39:24, April. 

= Microsciurus alfari browni (Bangs, 1902). 

See J. A. Allen (1914:151). 

Holotype. MCZ 10404. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Bogaba, 600 ft (183 
m). 15 July 1901. 



106 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Collector. W. W. Browii, Jr. Original number 631. 
Condition. Sldn and skull complete. 
Type Series. 4 paratypes; MCZ 10405, adult fe- 
male, 10406, adult female, 10407, adult female, 
10408, juvenile male; all represented by sldn and 
skull. 

Comments. M. a. broivni was retained as a valid 
subspecies by Hall (1981: 439). 

Genus PARAXERUS Forsyth Major, 1893 

Aethosciurus byatti laetus G. M. Allen and 

Loverldge, 1933 

Bull. Mus. Comp. Zool, 75: 96, 

Februaiy. 

= Paraxerus vexillarius byatti (Kershaw, 

1923). See Amtmann (1975: 11). 

Holotype. MCZ 26198. Skin and skull. Adult male. 
Locality. Tanganyika Territory (=Tanzania): north 
end of Lake Nyasa, Ukinga Mountains, Madehani, 
7,000 ft (2,135 m). 22 Februaiy 1930. 
Collector. A. Love ridge. 
Condition. Sldn and skull complete. 
Type Series. 9 paratypes; MCZ 26196, 26197, 
26199-26202, 26204-26206; all represented by 
skin and skull, 4 females and 5 males. 

Genus SCIUROTAMIAS M\\\er, 1901 

Sciurotamias davidanus thayeri G. M. 

Allen, 1912b 

Mem. Mus. Comp. Zool., 40: 231, 

August. 

= Sciurotamias davidianus consobrinus 

(Milne-Edwards, 1868). See Moore and 

Tate (1965: 308). 

Holotype. MCZ 8008. Skin and skull. Adult male. 
Locality. China: western Szechwan ( = Sichuan), 
Washan (=Wushan), 6,000 ft (1,830 m). 17 May 
1908. 

Collector W. R. Zappey. Original number 163. 
Condition. Sldn complete. Skull partial (basioccip- 
ital and left maxilla damaged). Mandible disartic- 
ulated. 

Type Series. Holotype only. 

Comments. G. M. Allen (1912b: 231) noted in the 
original description of thayeri that "unfortunately, 
tlie skull of the type was lost." The sldn and skull 
have been revmited subsequently. 

Genus SC/L/RL/S Linnaeus, 1758 

Sciurus (Guerlinguetus) aestuans 
cfiiriquensis Bangs, 1902b 
Bull. Mus. Comp. Zool., 39: 22, April. 
= Sciurus granatensis chiriquensis Bangs, 
1902. See Hershkovitz (1947: 7). 

Holotype. MCZ 10044. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Divala. 18 November 
1900. 



Collector. W. W. Brown, Jr. Original number 10. 
Condition. Skin and skull complete. 
Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. S. g. chiriquensis was retained as a val- 
id subspecies by Hall (1981: 437) and Nitikman 
(1985: 1). 

Sciurus carolinensis extimus Bangs, 1896j 
Proc. Biol. Soe. Washington, 10: 158, 28 
December. 

Holotype. B4519. Sldn and skull. Adult female. 

Locality. (United States): Florida, Dade County, 

Miami. 12 March 1895. 

Collector. L. Brownell. Original number 59. 

Condition. Skin complete (with bald spots on ven- 

trum). Skull complete. 

Type Series. 7 paratypes; B3406, B4517, B4518, 

(all represented by skin and skull), B4520-B4523 

(skins only); 4 females and 3 males. 

Comments. Retained as a valid subspecies by Hall 

(1981: 417) and Koprowskd (1994: 1). 

Sciurus carolinensis var. yucatanensis J. 

A. Allen, 1877 

In Coues and J. A. Allen, Monogr. N. 

Amer. Rodentia, U.S. Geol. Geograph. 

Suivey Terr., Rep. 11: 705, August. 

= Sciurus yucatanensis J. A. Allen, 1877. 

See Elliot (1896: 80). 

Syntype. MCZ 5398. Skin. Adult male. 

Locality. (Mexico): Yucatan, Merida. March 1865. 
Collector. A. Schott. Original number 228. For- 
merly USNM 8502. 
Condition. Sldn complete. 

Type Series. J. A. Allen referred to "four specimens 
of this variety before me" in the original descrip- 
tion (1877: 705). Three specimens are mentioned 
by number: USNM 8502 (now MCZ 5398), 8503, 
and 8505. A juvenile referred to in the description 
(but not by number) corresponds to USNM 8504. 
Comments. S. yucatanensis was considered a valid 
species by Hoffman et al. (1993: 443) and Nowak 
(1999: 1265). The syntype of yucatanensis in the 
MCZ was received from the USNM in March 
1877. Poole and Schantz (1942: 554) state that 
USNM 8505 is no longer in the USNM. 

Sciurus ludovicianus vicinus Bangs, 1896j 

Proc. Biol. Soc. Washington, 10: 150, 28 

December. 

= Sciurus niger vulpinus Gmelin, 1896. 

See Bark-alow (1954: 25). 

Holotype. B5215. Skin and skull. Adult female. 
Locality. (United States): West Virginia, Greenbri- 



Type Specimens of Recent Mammals • Helgen and McFodden 107 



ei" County, White Sulphur Springs. 29 Januaiy 

1896. 

Collector. T. Surber. Original number 55. 

Condition. Sldn and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Sciurus nesaeus G. M. Allen, 1902b 

Proc. Biol. Soc. Washington, 15: 93, 25 

April. 

= Sciurus granatensis nesaeus G. M. 

Allen, 1902. See Hershko\atz (1947: 37). 

Holotijpe. MCZ 10744. Skin. Adult female. 

Locality. Venezuela: (Nueva Esparta), Margarita 

Island, El Valle. 8 July 1901. 

Collector. A. H. Clark. Original number 619. 

Condition. Skin complete. The lower incisors are 

present in the sldn. 

Type Series. Holotype only. 

Comments. S. g. nesaeus was retained as a valid 

subspecies by Cabrera (1961: 367) and Nitikman 

(1985: 1). 

Sciurus variabilis morulus Bangs, 1 900d 

Proc. New England Zool. Club, 2: 43, 20 

September. 

= Sciurus granatensis morulus Bangs, 

1900. See Miller and Kellogg (1955: 

257). 

Holotype. B8420. Skin and skull. Adult female. 

Locality. Panama: Canal Zone, Loma del Leon. 13 

March 'l900. 

Collector W. W. Brown, Jr. Original number 5. 

Condition. Skin complete. Skull slightly damaged 

(left bulla broken). 

Type Series. 5 paratypes; B8418, B8419, B8421- 

B8423 (juvenile); all represented by skin and skull, 

3 females and 2 males. 

Comments. S. g. mondus was retained as a valid 

subspecies by Hall (1981: 437) and Nitikman 

(1985: 1). 

Sciurus variabilis saltuensis Bangs, 1898o 

Proc. Biol. Soc. \\'ashington, 12: 185, 16 

November. 

= Sciurus granatensis saltuensis Bangs, 

1898. See Hershkovitz (1947: 15). 

Holotype. B8144. Skin and skull. Adult female. 

Locality. Colombia: Magdalena, Santa Marta 

Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26 

March 1898. 

Collector. W. W. Brown, Jr. Original number 112. 

Condition. Sldn and skull complete. 

Type Series. 2 paratypes; B8145, skin and skull, 

adult male; B8244, skin and skull, adult female. 

Comments. S. g. saltuensis was retained as a valid 



subspecies by Cabrera (1961: 368) and Nitikman 

(1985: 1). 

Genus SPERMOPHILUS F. Cuvier, 1825 

Citellus obscurus siccus G. M. Allen, 1925 
Amer. Mns. Novitates, 163: 3, 2 April. 
= Spermophilus alashanicus Buchner, 
1888. See Hoffmann et al. (1993: 444). 

Holotype. MCZ 19924. Skin and skull. Adult female. 
Locality. China: Shansi ( = Slienxi), 10 miles (16.1 
km) west of Taiyuanfu. August 1921. 
Collector. F. R. Wulsin. Original number 146. 
Condition. Skin complete. Skull partial (parietals 
and basiooccipital missing, right tympanic bulla 
broken but present). Mandible disarticulated. 
Ti/pe Series. Holotype only. 

Spermophilus armatus Kennicott, 1863 
Proc. Acad. Nat. Sci. Philadelphia, 15: 
158, June. 

Sipitypes. MCZ 297: Skull. Male. Collected 11 April 
' 1858. Original number 167, formerly USNM 4799. 
MCZ 4793: Skin and skull. Male. Collected 2 April 
1858. Original number 140, formerly USNM 3478 
(3373). MCZ 4790: Skin. Female. Collected 26 
May 1858. Original number 455, formerly USNM 
3470 (3470). MCZ 4794: Skull only (skin spoiled in 
repreparation, discarded). Male. Collected April 
14, 1858. Original number 215, formerly USNM 
3474 (3373). 

Locality. (United States): Utah (now Wyoming), 
(Uinta County), near Fort Bridger, foothills of the 
Uinta Mountains, Camp Scott. 
Collector C. Drexler. 
Type Series. See comments below. 
Comments. Considered a valid species by Hoff- 
mann et al. (1993: 444) and Nowak (1999: 1254). 
Kennicott does not designate a type in the original 
descidption; thus, "all the specimens from Fort 
Bridger collected by C. Drexler and in the collec- 
tions [of the USNM] prior to 1863 are evidently 
cotypes [ = syntypes] of this species" (Lyon and Os- 
good 1909: 163). The syntypes of annatus in the 
MCZ were received from the USNM in January 
1874. These specimens possess USNM labels that 
bear numbers in discrepancy with the USNM cat- 
alogue; the numbers listed above are taken from 
the catalogue of the USNM, followed in parenthe- 
ses by the number on the specimen label. A com- 
plete list of syntypes, most of the remainder of 
which are in the USNM, can be found in Poole 
and Schantz (1942: 504-505). The skin of MCZ 
297, which formerly bore tlie number USNM 
3472, is not to be found in the collections of either 



108 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Spermophilus elegans Kennicott, 1863 
Proc. Acad. Nat. Sci. Philadelphia, 15: 
158, June. 

Syntijpes. MCZ 4791: Skin. Female. Collected 11 
April 1858. Original number 168. Formerly USNM 
3468. MCZ 4792: Skin. Male. Collected 11 April 
1858. Original number 166. Formerly USNM 
3473. 

Locality. (United States): Utah (now Wyoming), 
(Uinta County), Fort Bridger 
Collector. C. Dre.-der. 

Condition. Following their receipt, these speci- 
mens were spoiled in repreparation and discarded. 
Type Series. See comments below. 
Comments. These specimens are no longer in ex- 
istence. Kennicott does not designate a type in the 
original description; thus, "all the specimens from 
Fort Bridger collected by C. Drexler and in the 
collections [of the USNM] prior to 1863 are evi- 
dently cotypes [ = syntypes] of this species" (Lyon 
and Osgood 1909: 163). These syntypes oi elegans 
were received from the USNM in January 1874. A 
complete list of syntypes can be found in Lyon and 
Osgood (1909: 166). G. M. Allen (1931: 251) er- 
roneously included MCZ 4791 in a list of the type 
series of Spennophihis annatus. S. elegans is con- 
sidered a valid species by Hoffmann et al. (199.3: 
446) and Nowak (1999: 1254). 

Spermophilus (Ictidomys) tridecemlineatus 

badius Bangs, 1899c 

Proc. New England Zool. Club, 1: 1, 8 

Februaiy. 

= Spermophilus tridecemlineatus texensis 

Merriam, 1898. See Hall and Kelson 

(1959: 347). 

Holotype. B1682. Skin and skull. Adult male. 

Locality. (United States): Missouri (Vernon Coun- 
ty), Stotesbury. 17 April 1894. 
Collector. T. Surber. Original number 81. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 2 paratypes; B1683, skin and skull, 
adult male; B5609, skin and skull, adult male. 
Comments. A. H. Howell (1938: 110) first synon- 
yinized badius with texensis under die genus Ci- 
tellus. 

Genus SYNTHEOSCIURUS Bangs, 1902 

Syntheosciurus brochus Bangs, 1 902b 
Bull. Mus. Comp. Zool., 39: 25, April. 

Holotype. MCZ 10402. Sldn and skull. Adult male. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 7,000 ft (2.135 m). 30 April 1901. 
Collector W. W. Brown, Jr Original number 415. 



Condition. Skin complete. Skull partial (left supra- 
occipital damaged) 

Type Series. 1 paratype; MCZ 10403, sldn and 
skull, adult female. 

Comments. Type species of the genus Syntheo.sciu- 
nis Bangs, 1902. Considered a valid species by 
Hoffman et al. (1993: 452) and Nowak (1999: 
1268). Following a discussion with a collector em- 
ployed by W. W. Brown, Jr., and a biological survey 
of Boquete, Enders (1953b: 509) recommended 
that the type locality be considered "the Cordillera 
about 8 miles [12.9 km] north of Boquete and not 
at Boquete which is on the lower slopes of El Vol- 
can de Chiriqui." 

Genus LAMMS llliger, 1811 

Tamias cooperi Baird, 1855 

Proc. Acad. Nat. Sci. Philadelphia, 7: 

334, 24 April 24. 

= Tamias townsendii cooperi Baird, 1855. 

See Baird (1858: 737). 

Syntype. MCZ 4754. Skin only. Adult, unsexed. 
Locality. (United States): Washington (Skagit 
County), Clickitat ( = Klickitat) Pass, Cascade 
Mountains, 4,500 ft (1,373 m). July 1853. See com- 
ments. 

Collector J. G. Cooper. Formerly USNM 211/ 
1182. 

Condition. Sldn complete. 

Type Series. The other syntype is USNM 212/1183, 
skin and skull, unsexed adult. 

Comments. No type is designated in the original 
description, but Baird subsequently ex-plained that 
the two specimens listed above were those on 
which cooperi was based (1858: 301). Cooper 
(1869: .531) emended the type locality. The syntype 
of cooperi in the MCZ was received from the 
USNM in Januarv' 1874. 

Tamias dorsalis Baird, 1855 

Proc. Acad. Nat. Sci. Philadelphia, 7: 

332, 24 April. 

= Tamias dorsalis dorsalis Baird, 1855. 

See Hayman and Holt (1940: 435). 

Syntype. MCZ 4759. Skin and mandible. 

Locality. (United States): New Me.xico (Grant 
County), "Fort Webster, Coppermines of die Mim- 
bres" (near present Georgetown). 32°47'N, 
108°41'W. 1851. See A. H. Howell (1929: 131). 
Collector J. H. Clark. Formerly USNM 119/3151. 
Condition. Skin complete. Mandible partial (pos- 
terior of right mandibular ramus broken off and 
missing). 

Type Series. The other syntype is USNM 120, skull 
within skin. 

Comments. No type is designated in the original 
description, but Baird subsequently listed the two 



Type Specimens of Recent Mammals • Helaen and McFadden 



109 



specimens above as those on which dorsalis was 
based (1858: 300). The syntype of dorsalis in the 
MCZ was received from the USNM in January 
1874. 

Tamias quadrivittatus neglectus J. A. 

Allen, 1890 

Bull. Amer. Mus. Nat. Hist., 3: 106, 

June. 

= Tamias minimus neglectus (J. A. Allen, 

1890). See Hayman and Holt (1940: 

430). 

Holotype. MCZ 1575. Sldn and cranium. 

Locality. (Canada): (Ontario), eastern end of Lake 
Superior (near mouth of Montreal River). 5 July 
1848. 

Collector. L. Agassiz. 

Condition. Sldn complete. Cranium partial (zygo- 
matic arches, parietals, left tympanic bulla, and 
mandible missing). 

Type Series. The description mentions 6 paratypes, 
including MCZ 1567, skin and skull; and 4 others 
at USNM. 

Comments. Following A. H. Howell (1929: 54), 
Hayman and Holt (1940: 430) considered T. m. 
neglectus a synonym of T. m. borealis. However, 
neglectus was retained as a valid subspecies of min- 
imus by Hall (1981: 346) under the genus Euta- 
mias. 

Tamias striatus venustus Bangs, 1 896h 
Proc. Biol. Soc. Washington, 10: 137, 28 
December. 

Holotype. B5478. Skin and skull. Adult male. 

Locality. (United States): Indian Territory 

( = Oklahoma) (Adair County), Stilwell. 13 August 

1896, 

Collector T. Surber. Original number 63. 

Condition. Skin complete. Skull partial (condyle of 

left mandibular ramus inissing). 

Type Series. 2 paratypes; B5479, skin and skull, 

adult female; B5605, sldn and skull, adult male. 

Comments. Retained as a valid subspecies by Hall 

(1981: 340). 

Genus r>4/WMSC/L/f?L/S Trouessart, 1880 

Sciurus iiudsonicus gymnicus Bangs, 

18991 

Proc. New England Zool. Club, 1: 28, 31 

March. 

= Tamiasciurus Iiudsonicus gymnicus 

(Bangs, 1899). See Osgood (1938: 438). 

Holotype. B4914. Skin and skull. Adult female. 
Locality. (United States): Maine, Piscataquis 



County, Greenville, near Moosehead Lake. 1 De- 
cember 1895. 

Collector C. H. Goldthwaithe. Original number 2. 
Condition. Skin and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. T. h. gymnicus was retained as a valid 
subspecies by Hall (1981: 442) and Steele (1998: 
1). 

Sciurus hudsonicus loquax Bangs, 1 896j 

Proc. Biol. Soc. Washington, 10: 161, 28 

December. 

= Tamiasciurus hudsonicus loquax 

(Bangs, 1896). See A. H. Howell (1936: 

1). 

Holotype. B4270. Skin and skull. Adult male. 

Locality. (United States): Connecticut, New Lon- 
don County, Liberty Hill. 24 December 1895. 
Collector. O. Bangs. Original number 3. 
Condition. Skin and skull complete. 
Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. T. h. loquax was retained as a valid sub- 
species by Hall (1981: 443) and Steele (1998: 1). 

Sciurus hudsonicus orarius Bangs, 18971 

Proc. Biol. Soc. Washington, 11: 281, 30 

December. 

= Tamiasciurus douglasii mollipilosus 

(Audubon and Bachman, 1841). See 

Hayman and Holt (1940: 347). 

Holotype. B4978. Skin and skull. Adult female. 

Locality. (United States): California, Mendocino 
County, Philo. 9 December 1895. 
Collector. C. A. Allen. Original number 887. 
Condition. Skin complete. Skull partial (right tym- 
panic bulla broken). 

Type Series. 13 paratypes; B4832, B4979-B4989, 
B5462; all represented by sldn and skull, 9 feinales 
and 4 males. 

Family CASTORIDAE Hemprich, 1820 

Genus C/^SrOR Linnaeus, 1758 

Castor caecator Bangs, 1 91 3 
Bull. Mus. Comp. Zool., 54: 513, July. 
= Castor canadensis caecator Bangs, 
1913. See G. M. Allen (1942: 62). 

Holotype. B6979. Skull. Adult male. 

Locality. (Canada): Newfoundland, near Bay St. 
George. 1896. 
Collector. E. Doane. 

Condition. Skull complete. Mandible disarticulat- 
ed. 



110 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2 



Type Series. Holotype only. 

Comments. C. c. caecator was retained as a valid 

subspecies by Hall (1981: 602). 

Family GEOMYIDAE Bonaparte, 1845 

Genus GEOMVS Rafinesque, 1817 

Geomys colonus Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 178, 15 

March. 

= Geomys pinetis pinetis Rafinesque, 

1817. See Williams and Genoways (1980: 

444). 

Holotijpe. B5001. Skin and skull. Advilt male. 

Locality. (United States): Georgia, Camden Coun- 
ty, Arnot Plantation, about 4 miles (6.4 km) west 
of St. Marys. 21 March 1896, 
Collector. O. Bangs. Original number 8. 
Condition. Sldn and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. Hall (1981: 505) recognized Geomys 
colonus as a valid species. 

Geomys cumberlandius Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 180, 15 

March. 

= Geomys pinetis pinetis Rafinesque, 

1817. See Wihiams and Genoways (1980: 

444). 

Holotype. B5016. Skin and skull. Adult male. 

Locality. (United States): Georgia, Camden Coun- 
ty, Cumberland Island, Stafford Place. 17 April 
1896. 

Collector O. Bangs. Original number 1. 
Condition. Sldn and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. In the original description. Bangs lists 
B5015 as the type and gives its data. The data given 
lit B5016, not B5015. B5016 has "Type" written 
after the entiy in Bangs' catalogue and should be 
considered the holotype, rather than B5015. Hall 
(1981: 505) and Laerm (1981: 150) supported the 
specific status of Geomys cumherlandius. 

Geomys floridanus austrinus Bangs, 

1898b 

Proc. Boston Soc. Nat. Hist., 28: 177, 15 

March. 

= Geomys pinetis pinetis Rafinesque, 

1817. See WilHams and Genoways (1980: 

444). 

Holotype. B6983. Skin and skull. Adult male. 

Locality. (United States): Florida, Pinellas County, 
Belleair. 3 August 1897. 



Collector. W. S. Dickinson. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. 1 paratype, in the USNM. 

Geomys tuza gfof// Sherman, 1944 
Proc. New England Zool. Club, 23: 38, 
30 August. 

= Geomys pinetis goffi Slierman, 1944. 
See Harper (1952: 37). 

Holotype. B7222. Skin and skull. Adult male. 

Locality. (United States): Florida, Brevard County, 

Eau Gallie. 18 March 1897. 

Collector. O. Bangs. Original number 1. 

Condition. Sldn and skull complete. 

Type Series. 12 paratypes; B7212-B7217, B7219- 

B7221; all represented by sldn and skull, 5 females 

and 7 males. 

Comments. Williams and Genoways (1980: 444) 

synonymized gojfi with G. pinetis pinetis; however. 

Hall (1981: 504) maintained G, p. goffi as a valid 

subspecies. The lUCN designates goffi as extinct. 

Genus ORTHOGEOMYS Merriam, 1895 

IVIacrogeomys cavator Bangs, 1902b 
Bull. Mus. Conip. Zool., 39: 42, April. 
= Orttiogeomys cavator cavator (Bangs, 
1902). See Russell (1968a: 532). 

Holotijpe. MCZ 10381. Skin and sk-ull. Adult male. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 4,800 ft (1,464 m). 9 March 1901. 
Collector W. W. Brown, Jr. Original number 212. 
Condition. Skin and skull complete. Mandible dis- 
articvdated. 

Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. O. cavator was considered a valid spe- 
cies by Patton (1993: 470) and Nowak (1999: 1314). 

Macrogeomys pansa Bangs, 1902b 
Bull. Mus. Comp. Zool., 39: 44, April. 
= Ortiiogeomys cavator pansa (Bangs, 
1902). See Russell (1968a: 532). 

Holotype. MCZ 10364. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Bogaba, 600 ft (183 
m). 6 July 1901. 

Collector W. W. Brown, Jr Original number 567. 
Condition. Skin complete. Skull partial (right tym- 
panic bulla missing). 

Type Series. There are 7 paratypes; MCZ 10362— 
10363, 10365-10369, 4 females and 3 males, all 
represented by skin and skull. MCZ 10362 and 
10366 are now in the FMNH, and MCZ 10365 is 
in the USNM. 

Comments. O. c. pansa was retained as a valid sub- 
species by Hall (1981: 513). 



Type Specimens of Recent Mammals • Helaen and McFadden 111 



Orthogeomys grandis pluto Lawrence, 

1933a 

Proc. New England Zool. Club, 13: 66, 8 

May. 

Holotype. MCZ 29040. Skin and skull. Adult female. 
Locality. Honduras: Francisco Morazan, north of 
Tegucigalpa, Cerro Cantoral. 20 July 1932. 
Collector. C. F. Underwood. Original number 
1100. 

Condition. Skin and skull complete. 
Type Series. 2 paratypes; MCZ 29038, skin and 
sk-ull, juvenile female; MCZ 29039, skin and skull 
unsexed juvenile. 

Coninient.s. Retained as a valid subspecies by Hall 
(1981: 509). 

Genus PAPPOGEOMYS Mernam, 1895 

Cratogeomys castanops rubellus Nelson 

and Goldman, 1934a 

Proc. Biol. Soc. Washington, 47: 147, 13 

June. 

= Pappogeomys castanops rubellus 

(Nelson and Goldman, 1934). See Russell 

(1968b: 682). 

Holotype. MCZ 20507. Skin and skrdl. Adult male. 
Locality. Mexico: San Luis Potosi, near San Luis 
Potosi, Soledad, 6,400 ft (1,952 m). 1 August 1923. 
Collector W. W. Browai, Jr. Original number 2.31. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a large paratype series in the 
MCZ. 

Comments. P. a. nibellus was retained as a valid 
subspecies by Hall (1981: 520). 



Family HETEROMYIDAE Gray, 1868 

Genus DIPODOMYS Gray, 1841 

Dipodomys californicus pallidulus Bangs, 

1899J 

Proc. New England Zool. Club, 1: 65, 31 

July. 

= Dipodomys californicus californicus 

Merriam, 1890. See Kelt (1988: 1). 

Holotype. B9147. Skin and skull. Adult female. 
Locality. (United States): California, Colusa Coun- 
ty, Sites. 27 June 1896. 

Collector P. O. Simons. Original number 222. 
Condition. Skin complete. Skull partial (left jugal 
missing). 

Type Series. 1 paratype; B9148, sldn and skull, 
adult female. 

Dipodops ordii palmeri J. A. Allen, 1891 

Bull. Amer. Mus. Nat. Hist., 3: 276, 30 

June. 

= Dipodomys ordii palmeri {y}. A. Allen, 

1891). See Grinnell (1921: 96). 

Syntypes. MCZ 5886: Skin and skull. Juvenile male. 
MCZ 5887: Skin and skull. Unsexed juvenile. 
Locality. Mexico: San Luis Potosi, San Luis Potosi. 
1 May 1878. 
Collector E. Palmer. 

Condition. MCZ 5886: Sldn complete. Skull partial 
(part of parietal, right occipital condyle, and tym- 
panic bulla missing). MCZ 5887: Skin complete. 
Skull partial (left squamosal process missing). Man- 
dible disarticulated. 
Type Series. 2 syntypes only. 

Comments. Allen did not specify a holotype in the 
original description, which he based on the two 
specimens above. D. o. palmeri was retained as a 
vahd subspecies by Hall (1981: 569). 



Genus THOMOMYS Wied-Neuwied, 1839 Genus HETEROMYS Desmarest, 1817 

Heteromys repens Bangs, 1902b 
Bull. Mus. Comp. Zool, 39: 45, April. 
^Heteromys desmarestianus repens 
Bangs, 1902. See Goldman (1920: 115). 



Thomomys umbrinus atrodorsalis Nelson 

and Goldman, 1934b 

J. Mammal., 15: 111, 15 May. 

Holotype. MCZ 20487. Skin and skull. Adult male. 
Locality. Mexico: San Luis Potosi, Alvarez, 8,000 
ft (2,440 m). 7 November 1923. 
Collector W. W. Brown, Jr. Original number 338. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. Retained as a valid subspecies by Hall 
(1981: 476). 



Holotype. MCZ 10356. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 4,000 ft (1,220 m). 8 April 1901. 
Collector W. W. Brown, Jr. Original number 264. 
Condition. Sldn complete. Skull partial (left jugal 
and process of squamosal missing). 
Type Series. 5 paratypes; MCZ 10359, adult male, 
10355, adult male, 10358, adult male, 10361, ju- 
venile female, 10360, juvenile female. 
Comments. H. d. repens was retained as a valid 
subspecies by Hall (1981: 597). 



112 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Genus LIOMYS Uernam, 1902 

Heteromys alleni Coues, 1881 

In J. A. Allen, Bull. Mus. Comp. Zool., 8: 

187, March. 

= Liomys irroratus alleni (Coues, 1881). 

See Goldman (1911: 56). 

Holotype. MCZ 5889. Skin and skull Male. 

Locality. (Mexico): San Luis Potosi, Rio Verde, Ha- 
cienda Angostura. 26 Febniaiy 1878. 
Collector. E. Palmer. 

Condition. Sldn complete, skull withiji skin. 
Type Series. Holotype only. 

Comments. L. i. alleni was retained as a valid sub- 
species by Hall (1981: 590). 

Genus PEROGNATHUSW\e6-NeuW\e6, 
1839 

Perognathus longimembris bangs! Mearns, 

1898 

Bull. Amer. Mus. Nat. Hist., 10: 300, 31 

August. 

Holotype. B5304. Skin and skull. Adult female. 

Locality. (United States): California, (Riverside 
County), Colorado Desert, Palm Springs, 450 ft 
(137 m). 13 April 1896. 

Collector. E. C. Thurber. Original number 644. 
Condition. Sldn complete. Skull partial (teeth sep- 
arate from skull). Mandible disaiUculated. 
Type Series. 2 paratypes; B5302, adult male, skin 
and sloill; B5303, adult female, skin and skull. 
Comments. Retained as a valid subspecies by Hall 
(1981: 537). 

Family DIPODIDAE Fischer de Waldheim, 
1817 

Genus ZAPUS Coues, 1875 

Zapus hudsonius hardy! Batchelder, 1 899 

Proc. New England Zool. Club, 1: 5, 8 

February. 

=Zapus hudsonicus acadlcus (Dawson, 

1856). See Krutzsch (1954: 432). 

Holotype. MCZ 41681. Skin and skull. Adult female. 
Locality. (United States): Maine, Hancock County, 
Mount Desert Island. 24 August 1898. 
Collector C. F. Batchelder. Original number 1597. 
Condition. Skin and skull complete. 
Tijpe Series. There is a series of paratypes in tlie 
MCZ. 



Zapus hudsonius ladas Bangs, 1 899d 
Proc. New England Zool. Club, 1: 10, 28 
February. 

Holotype. B4169. Skin and skull. Adult female. 
Locality. (Canada): Labrador, Hamilton Inlet, Ri- 
goulette. 18 July 1895. 

Collector. C. H. Goldthwaite. Original number 2. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a large series of paratypes, 
most of which are in the MCZ. 
Comments. Retained as a valid subspecies by Hall 
(1981: 843). 

Zapus orahus Preble, 1899 
N. Amer. Fauna, 15: 29, 8 August. 
=Zapus trinotatus orarlus Preble, 1899. 
See Hooper (1944: 67). 

Holotype. B250. Skin and skull. Adult male. 

Locality. (United States): California, Marin Coun- 
ty, Point Reyes. 14 May 1893. 
Collector C. A. Allen. Original number 618. 
Condition. Skin and skull complete. 
Type Series. 3 paratypes, not in MCZ; mentioned 
by locality in original description. 
Comments. Z. t. orarius was retained as a valid sub- 
species by Hall (1981: 846) and Gannon (1988: 1). 

Family MURIDAE llliger, 1811 

Subfamily ARVICOLINAE Gray, 1821 

Genus CH/OA/O/V/yS Miller, 1908 

Hypudaeus nivlcola Schinz, 1845 

Syst. Verzeichniss Saugethiere Synopsis 

Mammalium, 2: 236. 

= Chionomys nivalis (Martins, 1842). See 

Musser and Carleton (1993: 507). 

Syntype. MCZ 1291. Skin. Juvenile female. 
Locality. Switzerland: St. Gotthard. 
Collector Received from L. Agassiz, 
Condition. Sldn complete. 

Type Series. Schinz wrote that he had e.xamined 16 
specimens, "junge und alte." 

Comments. G. M. Allen discussed the somewhat 
uncertain tyjie status of this specimen (1931: 263). 
The original label bears the following: "Hypodaeus 
[sic] nivicola juv. 9 sp. nov. du St. Gotthard et du 
Faulhorn." Louis Agassiz, a student of Schinz's, de- 
posited the specimen in the MCZ. 



Type Specimens of Recent Mammals • Helgen and McFadden 113 



Genus CLETHRIONOMYSJWes'wJs, 1850 Genus EOTHENOMYS Miller, 1896 



Evotomys proteus Bangs, 1 897g 

In Bailey, Proc. Biol. Soc. Washington, 

11: 137, 13 May. 

= Clethrionomys gapperi proteus (Bangs, 

1897). See Jackson (1938: 433). 

Holotype. B4081. Sldn and skull. Adult female. 

Locality. (Canada): Labrador, Hamilton Inlet. 27 
August 1895. 

Collector. C. H. Goldthwaite. Original number 10. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a large series of paratypes in 
tlie MCZ. 

Comments. C. g. proteus was retained as a valid 
subspecies by Hall (1981: 783). 

Genus DICROSTONYX Gloger, 1841 

Dicrostonyx chionopaes G. M. Allen, 

1914b 

Proc. New England Zool. Club, 5: 62, 9 

April. 

= Dicrostonyx torquatus chionopaes G. M. 

Allen, 1914. See Ognev (1948: 507). 

Holotype. MCZ 15263. Skin and skull. Adult male. 
Locality. U.S.S.R. ( = Russian Federation): eastern 
Siberia, Nijni Kolymsk (Nizhnekolymsk), near 
mouth of Kolyma River. 15 October 1911. 
Collector J. Koren. Original number 257. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Dicrostonyx exsuKj. M. Allen, 1919a 
Bull. Mus. Comp. Zool., 62: 532, 
February. 

Holotype. MCZ 11885. Skin and skull. Adult male. 
Locality. (United States): (Alaska), St. Lawi-ence Is- 
land. 24 June 1913. 

Collector J. Dixon. Original number 3267. 
Condition. Skin complete. Skull partial (basioccip- 
ital and right tympanic bulla missing). Mandible 
disarticulated. 

Type Series. 3 paratypes; MCZ 11883, skin and 
skull, female; MCZ 11884, skin and skull, female; 
USNM 232007, skin and skull, female. 
Comments. Considered a valid species by Musser 
and Carleton (1993: 510) but included in D. groen- 
landicus by Nov^^ak (1999: 1479). 



Craseomys aquilus G. M. Allen, 1912b 

Mem. Mus. Comp. Zool, 40: 216, 

August. 

= Eotlienomys eva eva (Thomas, 1911). 

See G. M. Allen (1940: 837). 

Holotype. MCZ 7190. Skin and skull. Adult male. 
Locality. China: Hupeh ( = Hubei), Showlungtan. 
17 May 1907. 

Collector W. R. Zappey Original number 10. 
Condition. Sldn complete. Skull partial (hole in left 
parietal, occiput missing). 

Type Series. 5 paratypes; MCZ 7189, 7191-7194, 
7196; all represented by sldn and skull, all female. 

Microtus (Eothenomys) aurora G. M. 

Allen, 1912b 

Mem. Mus. Comp. Zool., 40: 211, 

August. 

= Eotlienomys melanogaster aurora (G. M. 

Allen, 1912). See Hinton (1923: 149). 

Holotype. MCZ 7788. Skin and skull. Male. 

Locality. China: Hupeh ( = Hubei), Changyangh- 

sieh. 2 Februaiy 1909. 

Collector W. R. Zappey. Original number 372. 

Condition. Skin and skull complete. 

Type Series. 3 paratypes; MCZ 7185, sldn and skull, 

female; MCZ 7186, skin and skull, male; MCZ 

7188, skin and skull, male. 

Comments. E. ni. aurora was retained as a valid 

subspecies by Zhang et al. (1997: 229). 

Microtus (Eotlienomys) mucronatus G. M. 

Allen, 1912b 

Mem. Mus. Comp. Zool., 40: 214, 

August. 

= Eotiienomys melanogaster melanogaster 

(Milne-Edwards, 1871). See G. M. Allen 

(1940: 806). 

Holotype. MCZ 7789. Sldn and skull. Aduk female. 
Locality. China: western Szechwan ( = Sichuan), 
Tachiao, 12,000 ft (3,660 m). 11 August 1908. 
Collector. W. R. Zappey. Original number 269. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; MCZ 7790, sldn and skull, 
adult female (exchanged to the BMNH); MCZ 
7791, skin and skuU, adult female; MCZ 7803, skin 
and skuU, juvenile female. 



114 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Genus LEMMUS Link, 1795 

Lemmus paulus G. M. Allen, 1914b 
Proc. New England Zool. Club, 5: 60, 9 
April. 

= Lemmus sibiricus chrysogasterJ. A. 
Allen, 1903. See Ellerman and Morrison- 
Scott (1951: 656). 

Holotijpe. MCZ 15268. Skin and skull. Adult male. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Kalascliowo, near mouth of Kolyma River. 22 
June 1912. 

Collector. J. Koren. Original number 152. 
Condition. Skin complete. Skull partial (left man- 
dibular ramus missing). 
Ti/pe Series. 1 paratype; MCZ 15267, skin, female. 

Genus M IC ROTUS Schrank, 1798 

Arvicola breweri Bah6, 1858 
Mammals, in Repts. Explor. Surveys 
Railr. to Pacific, 8(1): 525, 14 July. 
= Microtus breweri {Bai'cd, 1858). See 

Miller (1896: 83). 

Syntype. MCZ 4365. Body in alcohol. Formerly 
USNM 2833. 

Locality. (United States); Massachusetts, Nantuck- 
et County, off Nantucket Island, Muskeeget ( = Mu- 
skeget) Island. July 1856. 
Collector. T. M. Brewer. 
Condition. Alcoholic. 

Type Series. There are 5 other syntypes (see Poole 
and Schantz, 1942: 271-272), all of which were at 
one time in the USNM. 

Comments. Poole and Schantz (1942: 271) noted 
that USNM 2829, a svntype of breweri, could not 
be found. 

Arvicola riparia var. longipilis Baird, 1858 
Mammals, in Repts. Explor. Sin-vevs 
Railr. to Pacific, 8(1): 524, 14 July' 
= Microtus pennsylvanicus pennsylvanicus 
(Ord, 1815). See Bailey (1900: 16). 

Syntype. MCZ 5292. Skin and skull. 

Locality. (United States): Illinois, (Cook County), 
West Northfield. Spring 1855. 
Collector R. Kennicott. Formerly USNM 745. 
Condition. Sldn complete. Skull partial (part of pa- 
rietal, both tympanic bullae and palatine missing). 
Mandible disarticulated. 
Tijpe Series. See comments. 

Comments. G. M. Allen (1931: 260), mentioned 
that "although no individuals are mentioned [in 
Baird's description], . . . [tliis] specimen was un- 
doubtedly among those examined by Baird in pre- 
paring his diagnosis and, therefore, is a cotype 



[ = syntype]." Baird's specimens were collected by 
Kennicott at West Northfield, Illinois, and by Hoy 
at Racine, Wisconsin, and were apparently depos- 
ited in the USNM. Lyon and Osgood (1909) and 
Poole and Schantz (1942) make no mention of oth- 
er sxii types. 

Arvicola rufidorsum Baird, 1858 

Repts. Explor. Surveys Railr. to Pacific, 

8(1): 526, 14 July. 

= Microtus pennsylvanicus pennsylvanicus 

(Ord, 1815). See Bailey (1900: 16). 

Holotype. MCZ 54372. Skin and skull. 

Locality. (United States): Massachusetts, (Duke's 
County), Martha's Vineyard, Holmes Hole. 
Collector D. J. Wyman. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. This specimen has formerly been 
BSNH 1949 and USNM 901. 

Arvicola terraenovae Bangs, 1894a 

Proc. Biol. Soc. Washington, 9: 129, 27 

July. 

= Microtus pennsylvanicus terraenovae 

(Bangs, 1894). See Davis (1936: 290). 

Holotype. B1104. Skin and skull. Adult male. 

Locality. (Canada): Newfoundland, Codroy. 27 No- 
vember 1893. 

Collector E. Doane. Original number 4. 
Condition. Sldn and sk-ull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. M. p. terraenovae was retained as a val- 
id subspecies bv Hall (1981: 796). 

Microtus chrotorrhinus ravus Bangs, 

1898p 

Proc. Biol. Soc. Washington, 12: 188, 16 

November. 

Holotype. B7951. Skin and skull. Adult male. 

Locality. (Canada): Labrador, Strait of Belle Isle, 
Black Bay. 15 July 1898. 
Collector. E. Doane. Original number 4. 
Condition. Skin and skull coiuplete. Mandible dis- 
aiticulated. 

Type Series. Paratype mateiial consists of Bangs' 
series from the type locahty; B7952-7967, all rep- 
resented by skin and skull, 10 females, 3 males, and 
3 unsexed individuals. B7953 and B7955 v^ere ex- 
changed to MVZ in 1937; B7964 was exchanged to 
USNM in 1922. 

Comments. Retained as a valid subspecies by Hall 
(1981: 811) and Kirkland and Jannett (1982:' 1). 



Type Specimens of Recent Mammals • Helaen and McFadden 



115 



Microtus enixus Bangs, 1896g 
Amer. Nat., 30: 1051, 1 December. 
= Microtus pennsylvanicus enixus Bangs, 
1896. See Davis (1936: 290). 

Holotype. B3973. Skin and skull. Adult female. 
Locality. (Canada): Labrador, Hamilton Inlet. 15 
July 1895. 

Collector. C. H. Goldthwaite. Original number 4. 
Condition. Skin and skull complete. 
Type Series. There is a large series of parat\'j3es in 
the MCZ. 

Comments. M. p. enixus was retained as a valid 
subspecies by Hall (1981: 79.3). 

Microtus fontigenus Bangs, 1896d 

Proc. Biol. Soc. Washington, 10: 48, 9 

March. 

= Microtus pennsylvanicus fontigenus 

Bangs, 1896. See Miller (1897: 14). 

Holotype. B3837. Skin and skull. Adult female. 

Localitij. (Canada): Quebec, Lake Edward. 28 Sep- 
tember 1895. 

Collector. E. A. and O. Bangs. Original number 9. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Tijpe Series. 7 paratypes; B3838-B3844; all repre- 
sented by sldn and skull, 2 females and 5 males. 
Comments. M. p. fontigenus was retained as a valid 
subspecies by Hall (1981: 793). 

Microtus l<oreni G. M. Allen, 1914b 

Proc. New England Zool. Club, 5: 64, 9 

April. 

= Microtus oeconomus koreni G. M. Allen, 

1914. See Ellerman and Morrison-Scott 

(1951: 706). 

Holotype. MCZ 15213. Skin and skull. Adult female. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth 
of Kolyma River. 1 November 1911. 
Collector J. Koren. Original number 132. 
Condition. Skin complete. Skull partial (left tym- 
panic bulla broken, right mandibular ramus miss- 
ing). 

Type Series. There is a large series of paratypes in 
the MCZ. 

Microtus pennsilvanicus [sic] sliattucl<i 

Howe, 1901 

Proc. Portland Soc. Nat. Hist., 2: 201, 31 

December. 

= Microtus pennsylvanicus shattucki 

Howe, 1901. See Wyman (1922: 162). 

Holotype. MCZ 10011. Sldn and skull. Adult female. 
Locality. (United States): Maine, Penobscot Bay, 



near Long Island, Tvmible Down Dick Island. 10 
July 1900. 

Collector R. H. Howe and G. C. Shattuck. Original 
number 31. 

Condition. Skin and skull complete. 
Type Series. There is a small series of paratypes in 
the MCZ. 

Comments. Wyman (1922: 166) considered M. p. 
shattucki a synonym of M. p. pennsylvanicus, but 
Hall (1981: 796) retained it as a separate subspe- 
cies. 

Microtus pennsylvanicus acadicus Bangs, 
1897c 

Amer. Nat., 31: 239, 1 March. 

Holotype. B2155. Skin and skull. Adult female. 

Locality. Canada: Nova Scotia, Digbv. 22 |uly 

1894. 

Collector O. Bangs. 

Condition. Sldn and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Comments. Retained as a valid subspecies by Hall 

(1981: 792). 

Microtus provectus Bangs, 1 908 

Proc. New England Zool. Club, 4: 20, 6 

March. 

= Microtus pennsylvanicus provectus 

Bangs, 1908. See Chamberlain (1954: 

589). 

Holotype. B9794. Skin and skull. Adult female. 

Locality. (United States): Rhode Island, Block Is- 
land. 5 August 1899. 

Collector O. Bangs. Original number 11. 
Condition. Sldn complete. Skull partial (tympanic 
bullae broken). Mandible disarticulated and angu- 
lar process chipped. 

Type Series. There is a series of paratypes in the 
MCZ. 

Comments. M. p. provectus was retained as a valid 
subspecies by Hall (1981: 796). 

Genus MYOPUSmWer, 1910 

Myopus thayeri G. M. Allen, 1914b 

Proc. New England Zool. Club, 5: 58, 9 

April. 

^Myopus schisticolor thayeri G. M. Allen, 

1914. See Ognev (1948: 526). 

Holotype. MCZ 15264. Sldn and skull. Adult male. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Yakutsk, Nijni Kolyinsk ( = Nizhnekolyinsk), 
near mouth of Kolyma River 28 March 1912. 
Collector J. Koren. Original number 264. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 



116 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Comment.^. 1 paratype; MCZ 15265, skin and skull, 
male. 

Genus ONDATRA Link, 1795 



Fiber obscurus Bangs, 1894b 

Proc. Biol. Soc. Washington, 9: 133, 15 

September. 

= Ondatra zibethicus obscurus (Bangs, 

1894). See Cameron (1959: 85). 

Holotype. B1155. Skin and skull. Adult female. 

Locality. (Canada): Newfoundland, Codroy. 14 
May 1894. 

Collector. E. Doane. Original number 3. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a small series of paratypes in 
the MCZ. 

Comments. O. z. obscurus was retained as a valid 
subspecies by Hall (1981: 827). 

Fiber zibethicus aquilonius Bangs, 1899d 

Proc. New England Zool. Club, 1: 11, 28 

February. 

= Ondatra zibettiicus aquilonius (Bangs, 

1899). See Miller (1912:231). 

Holotype. B3957. Skin and skull. Adult male. 

Locality. (Canada): Labrador, Hamilton Inlet, Ri- 
goulette. 15 August 1895. 

Collector. C. H. Goldthwaite. Original number 11. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; B7974, sldn and skull, 
adult female; B7975, sldn and skull, juvenile; 
B8704, skull, juvenile. 

Comments. O. z. aquilonius was retained as a valid 
subspecies by Hall (1981: 825). 

Fiber zibethicus rivalicius Bangs, 1895b 
Proc. Boston Soc. Nat. Hist., 26: 541, 31 

= Ondatra zibethicus rivalicius (Bangs, 
1895). See Davis and Loweiy (1940: 

212). 

Holotype. B2719. Sldn and skull. Adult male. 

Locality. (Canada): Louisiana, Plaquemines Parish, 

Burbridge. 31 January 1895. 

Collector. F. L. Small. Original number 1556/165. 

Condition. Sldn and skull complete. 

Type Series. 4 paratypes; B2720, skin and skull, 

adult female; B2721, skin and skull, adult male; 

B2883, skin and skull, male; B2884, sldn and skull, 

male. 

Comments. O. z. rivalicius was retained as a valid 

subspecies by Hall (1981: 827). 



Genus PHENACOMYS Mernam, 1889 

Phenacomys celatus crassus Bangs, 

1900c 

Proc. New England Zool. Club, 2: 39, 20 

September. 

= Phenacomys ungava crassus Bangs, 

1900. See A. B. Howell (1926: 27). 

Holotype. B3959. Skin and skull. Adult male. 

Locality. (Canada): Labrador, Hamilton Inlet, Ri- 

goulette. 15 August 1895. 

Collector C. H. Goldthwaite. Original number 1. 

Condition. Sldn and skull complete. 

Type Series. There is a small series of paratypes, 

most of which are no longer in the MCZ. 

Comments. In the original description, "3946" is a 

misprint for 3964. 

Genus SYNAPTOMYS Ba\r6, 1858 

Synaptomys fatuus Bangs, 1 896d 

Proc. Biol. Soc. Washington, 10: 47, 9 

March. 

= Synaptomys cooperi cooperi Baird, 

1858. See Wetzel (1955:8). 

Holotype. B3857. Skin and skull. Adult female. 
Locality. (Canada): Quebec, Lake Edward. 28 Sep- 
tember 1895. 

Collector E. A. and O. Bangs. Original number 3. 
Condition. Sldn and skull complete. 
Type Series. 8 paratypes; B3854-B3856, B3858- 
B3862; all represented by sldn and skull, 3 females 
and 5 males. 

Synaptomys (Mictomys) innuitus 

medioximus Bangs, 1 900c 

Proc. New England Zool. Club, 2: 40, 20 

September. 

= Synaptomys borealis medioximus 

Bangs, 1900. See A. B. Howell (1927: 9). 

Holotype. B8852. Skin and skull. Adult male. 

Locality. (Canada): Labrador, Strait of Belle Isle, 
Lance ( = Lanse) an Loup. 15 April 1899. 
Collector E. Doane. Original number 7. 
Condition. Sldn complete. Skull slightly damaged 
(hole in left parietal). 

Type Series. 1 paratype; B3972, sldn and skull, 
adult male. 

Comments. S. b. medioximus was retained as a val- 
id subspecies by Hall (1981: 834). 



Type Specimens of Recent Mammals • Helaen and McFadden 117 



Subfamily CRICETINAE Fischer de 
Waldheim, 1817 

Genus CANSUMYS G. M. Allen, 1928 

Cansumys canus G. M. Allen, 1928 
J. Mammal., 9: 245, 9 August. 

Holotype. MCZ 23779. Skin and skull. Adult female. 
Locality. China: southern Kansu (=Xinjiang), 
Choni. 9 December 1925. 
Collector. R. B. Ekvall. Original number 79. 
Condition. Skin complete. Skull partial (left jugal 
and occiput missing). 

Type Series. 1 paratype; MCZ 23780, skin and 
skull, juvenile male. 

Comments. Type species o( Cansumys G. M. Allen, 
1928. Considered a valid species by Musser and 
Carleton (1993: 537) and Nowak (1999: 1423). 
Known only by the type series and one additional 
specimen, FMNH 36067. 

Subfamily CRICETOMYINAE Roberts, 
1951 

Genus SACCOSTOMUS Peters, 1846 

Saccostomus cricetulus G. M. Allen and 
Lawrence, 1936 

Bull. Mus. Comp. Zool., 79: 100, January. 
= Saccostomus mearnsiHeWer , 1910. See 
Hubert (1978: 51). 

Holotype. MCZ 31475. Skin and skaill. Subadult male. 
Locality. Uganda: Sabei District, due north of 
Mount Elgon, south bank of Greek River ( = Kelim 
River), 3,000 ft (915 m). 5 December 1933. 
Collector. A. Loveridge. 

Condition. Skin complete. Skull partial (right tym- 
panic bulla missing). Mandible disarticulated. 
Type Series. 1 paratype; MCZ 31474, skin and 
skull, adult female. 

Subfamily DENDROMURINAE G. M. 
Allen, 1939 

Genus S7E/\ rO/WVS Peters, 1846 

Steatomys pratensis nyasae Lawrence 

and Loveridge, 1953 

Bull. Mus. Comp. Zool, 110: 39, June. 

Holotype. MCZ 44213. Skin and skull. Adult male. 
Locality. Nyasaland ( = Malawi): foot of Mulanje 
Mountain, Likabula River, 2,100 ft (641 m). 29 July 
1948. 

Collector. A. Loveridge. 
Condition. Sldn and skull complete. 
Type Series. 14 paratypes; MCZ 44214-44216, 
44218-44228; all represented by skin and skull, 9 
males and 5 females. 



Comments. Retained as a valid subspecies by An- 
sell (1978: 77). 

Subfamily GERBILLINAE Gray, 1825 

Genus TATERA Lataste, 1882 

Tatera flavipes G. M. Allen, 1914d 
Bull. Mus. Comp. Zool., 5S: 331, July. 
= Tatera valida /cemp/ Wroughton, 1906. 
See Bates (1988: 277). 

Holotype. MCZ 14491. Skin and skull. Adult female. 
Locality. Sudan: Blue Nile, north of (Er) Roseires, 
Aradeiba. 22 Januaiy 1913. 

Collector G. M. Allen and J. C. PhilHps. Original 
number 69. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Tatera sororG. M. Allen, 191 4d 
Bull. Mus. Comp. Zool, 58: 333, July 
= Tatera valida /cemp/ Wroughton, 1906. 
See Bates (1988: 277). 

Holotype. MCZ 14492. Skin and skull. Adult female. 
Locality. Sudan: Blue Nile, Fazogli. 16 January 
1913. 

Collector CM. Allen and J. C. Phillips. Original 
number 53. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; MCZ 14493, skin and 
skull, juvenile male. 

Genus TATERILLUS Thomas, 1910 

Taterillus melanops G. M. Allen, 1912a 
Bull. Mus. Comp. Zool., 54: 446, April. 
= Taterillus harringtoni (Thomas, 1906). 
See Musser and Carleton (1993: 563). 

Holotype. MCZ 8132. Skin and skull. Male. 

Locality. British East Africa ( = Kenya): arid plains 
by the Meru River. 11 August 1909. 
Collector G. M. Allen. Original number 108. 
Condition. Skin and skull complete. 
Tijpe Series. Holotype only. 

Subfamily MURINAE llliger, 1811 

Genus APODEMUS Kaup, 1829 

Apodemus mystacinus euxinus G. M. 

Allen, 1915b 

Bull. Mus. Comp. Zool, 59: 11, 

Februaiy. 

= Apodemus mystacinus mystacinus 

(Danford and Alston, 1877). See Corbet 

(1978: 133). 

Holotype. MCZ 14887. Skin and skull. Male. 

Locality. Asia Minor (=Turkey): (Trabzon), near 



118 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Trebizond (=Trabzon), Scalita, 1,000 m. 25 No- 
vember 1905. 

Collector. A. Robert. Original number 2189. 
Condition. Skin complete. Skull partial (right and 
left jugal missing, right tympanic bulla separated 
but present, right and left squamosal missing). 
Mandible disarticulated. 
Tijpe Series. HolotN-pe only. 

Genus DASYMYS Peters, 1875 

Dasymys incomtus alleni Lawrence and 

Loveridge, 1953 

Bull. Mus. Comp. ZooL, 110: 53, June. 

Holotijpe. MCZ 26322. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (=Tanzania): 
(Mbeya), Ilolo, near Rungwe Mountains, 4,600 ft 
(1,403 m). 31 March 1930. 
Collector. A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated and right coronoid process chipped. 
Type Series. 11 paratypes, all represented by sldn 
and skull, in the MCZ. 

Comments. Retained as a valid subspecies by An- 
sell (197S: S3). 

Genus GRAMMOMYS Jhomas, 1915 

Thamnomys ochraceus G. M. Allen, 

1912a 

Bull. Mus. Comp. ZooL, 54: 442, April 

1912. 

= Grammomys macmillani (Wroughton, 

1907). See Musser and Carleton (1993: 

594). 

Holotype. MCZ 8126. Skin and skull. Adult male. 
Locality. British East Africa ( = Kenya): Meru Riv- 
er, near junction with northern Guaso Nyiro 
( = Ewaso Ngiro). 8 August 1909. 
Collector. G. M. Allen. Original number 103. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Genus HyLO/WySCL/S Thomas, 1926 

Hylomyscus alleni simus G. M. Allen and 

Coolidge, 1930 

7/7 Strong, The African Republic of 

Liberia and the Belgian Congo, 2: 599, 

October. 

= Hylomyscus alleni (Waterhouse, 1838). 

See Heim de Balsac and Aellen (1965: 

722). 

Holotype. MCZ 24028. Skin and skull. Adult male. 
Locality. Liberia: Merikay. 13 September 1926. 



Collector G. M. Allen and H. J. Coolidge, Jr. Orig- 
inal number 97. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a series of paratypes in the 
MCZ. 

Genus MELO/W/S Thomas, 1922 

Melomys levipes stevensi Rummler, 1935 

Z. Saugetierkunde, 10: 109, 31 

December. 

= Melomys mollis Thomas, 1913. See 

Flanneiy (1990: 226). 

Holotype. MCZ 29890. Skin and skull. Adult male. 
Locality. Papua New Guinea: Morobe, Mount Mis- 
im ( = Missim), 6,700 ft (2,044 m). 17 April 1933. 
Collector H. Stevens. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 11 paratypes; MCZ 29889, 29892- 
29899, 29901; all represented by sldn and skull, 4 
females and 7 males. 

Melomys moncktoni alleni Rummler, 1935 

Z. Saugetierkunde, 10: 112, 31 

December. 

= Melomys rubex alleni Rummler, 1935. 

See Tate (1951: 299). 

Holotype. MCZ 29902. Skin and skull. Adult female. 
Localiti/. Papua New Guinea: Morobe, Mount Mis- 
im ( = Missim), 6,700 ft (2,044 m). 17 April 1933. 
Collector H. Stevens. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratvpe; MCZ 29900, skin and 
skull, adult male. 

Comments. Flannery (1995a: 308—309) did not list 
subspecies of M. nibex, noting, "It is a veiy variable 
species, and a taxonomic revision is needed in or- 
der to clarify the taxonomic status of many popu- 
lations." 

Genus MUS Linnaeus, 1758 

Leggada bufo ablutus G. M. Allen and 

Loveridge, 1942 

Bull. Mus. Comp. ZooL, 89: 199, 

Februaiy. 

= Mus bufo (Thomas, 1906). See Musser 

and Carleton (1993: 622). 

Holotype. MCZ 40745. Skin and skull. Adult male. 
Locality. Belgian Congo ( = Democratic Republic 
of Congo): Kivu, Lake Kivu, Idj\\d Island, Upper 
Mulinga, 6,500 ft (1,983 m). 24 February 1939. 
Collector A. Loveridge. 



Type Specimens of Recent Mammals • Helgen and McFadden 119 



Condition. Skin and skull complete. Mandible dis- 
articulated. 

Tijpe Series. 2 paratypes; MCZ 40746, skin and 
skull (no longer in MCZ); MCZ 40747, skin and 
skull, female. 

Leggada gerbillus G. M. Allen and 

Loveridge, 1933 

Bull. Mus. Comp. ZooL, 75: 112, 

Febriiaiy. 

= Mus tenellus (Thomas, 1903). See 

Musser and Carleton (1993: 629). 

Holotype. MCZ 26586. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (= Tanzania): Do- 
doma, Ugogo, 3,700 ft (1,129 m). 23 December 
1929. 

Collector. A. Loveridge. 

Condition. Sldn complete. Skull partial (right jugal 
missing). Mandible disarticulated and angular pro- 
cess of right ramus chipped. 
Type Series. Holotype only. 

Mus bactrianus tantillus G. M. Allen, 1927 
Amer. Mus. Novitates 270: 9, 31 May. 
= Mus musculus tantillus G. M. Allen, 
1927. See Marshall (1977: 214). 

Holotype. MCZ 23476. Skin and skull. Adult female. 
Locality. China: Sze-chuan ( = Sichuan), Wanhsien 
(=Wanxian). 14 November 1921. 
Collector W. W. Granger. 

Condition. Skin complete. Skull partial (6 frag- 
ments). Mandible disarticulated. 
Type Series. There is a series of paratypes in the 
AMNH, and 1 paratype in the MCZ; MCZ 23475 
(formerly AMNH 56403), skin and skull, male. 
Comments. This specimen was formerly AMNH 
56416. The original description lists AMNH 56413 
as the holotype, but this specimen is a juvenile, not 
an adult female, as Allen stated. The collection date 
and measurements of the holotype in the original 
description fit MCZ 23476 rather than any other 
specimens in the type series at the AMNH, which 
are all immatures (Lawrence 1993: 136). 

Genus /W/O/W/S Thomas, 1915 

Praomys fumatus oweni Setzer, 1 956 

Proc. U.S. Nat. Mus., 106: 525, 28 

November. 

= Myomys fumatus (Peters, 1878). See 

Musser and Carleton (1993: 631). 

Holotype. MCZ 45883. Skin and skull. Adult male. 
Locality. Anglo-Egyptian Sudan ( = Sudan): Equa- 
toria Province, Torit District, Murukurun, 50 miles 
(80.5 km) east of Torit, 2,000 ft (610 m). 9 May 
1950. 



Collector J. S. Owen. Original number 1030. 

Condition. Skin and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Genus A//l//\/EA/rE/? Marshall, 1976 

Epimys zappeyi G. M. Allen, 1912b 

Mem. Mus. Comp. ZooL, 40: 225, 

August. 

^Niviventer confucianus (Milne-Edwards, 

1871). See Musser and Carleton (1993: 

633). 

Holotype. MCZ 7607. Skin and skull. Adult male. 
Locality. China: western Szechwan ( = Sichuan), 
Washan Mountains (=Wu Shan), 9,000 ft (2,745 
m). 26 October 1908. 

Collector W. R. Zappey. Original number 305. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Genus OEA/OM/S Thomas, 1904 

Oenomys hypoxanthus talangae Setzer, 

1956 

Proc. U.S. Nat. Mus., 106: 505, 28 

November. 

Holotype. MCZ 45315. Skin and skull. Adult female. 
Locality. Anglo- Egyptian Sudan ( = Sudan): Equa- 
toria Province, Imatong Mountains, Talanga Foi- 
est, 3,000 ft (915 m). 10 July 1950. 
Collector J. S. Owen. Original number 1338. 
Condition. Skin and skull complete. 
Type Series. 1 parats-pe; MCZ 45284, skin and 
skuU, male. 



Genus PR/A 0/WVS Thomas, 1915 



Allen 



Praomys tullbergi melanotus G. 

and Loveridge, 1933 

Bull. Mus. Comp. ZooL, 75: 106, 

February. 

= Praomys delectorum (Thomas, 1910). 

See Musser and Carleton (1993: 642). 

Holotype. MCZ 26287. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (=Tanzania): north- 
west end of Lake Nyasa, Poroto Mountains, Nyam- 
wanga, 6,400 ft (1,952 m). 21 March 1930. 
Collector. A. Loveridge. 

Condition. Skin complete. Skull partial (left tym- 
panic bulla missing). Mandible disarticulated. 
Type Series. 23 paratypes; MCZ 26259, 26285, 
26286, 26288-26293, 26295-26297, 26387-26394, 
26411, 26497, 26498; all represented by skin and 
skull. 



120 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2 



Comments. Van der Straeten and Dieterlen (1987: 
9) and Van der Straeten and Dudu (1990: 81) treat- 
ed melanotus as a species within the Praomijs de- 
lectorum species complex. 

Genus PSEUDOHYDROMYS Rummler, 
1934 

Pseudohydromys murinus Rummler, 1 934 
Z. Saugetierkunde, 9: 48, 30 December. 

Holotype. MCZ 29904. Sldn and skull. Adult male. 
Locality. Papua New Guinea, Morobe, Mount Mis- 
im ( = Missim), 7,000 ft (2,135 m). 8 March 1933. 
Collector. H. Stevens. 

Condition. Sldn complete, with small bald spots. 
Skull complete. Mandible disarticulated. 
Type Series. Holotype only. 

Comments. Type species of Pseudohydromys Rii- 
mmler, 1934. Considered a valid species by Flan- 
nery (1995a: 255), Musser and Carleton (1993: 
644), and Nowak (1999: 1612). 

Genus STENOMYS Ihomas, 1910 

Stenomys niobe stevensi Rummler, 1935 

Z. Saugetierkunde, 10: 117, 31 

December. 

= Stenomys niobe niobe (Thomas, 1906). 

See Flamieiy (1995a: 339). 

Holotype. MCZ 29915. Skin and skull. Adult male. 
Locality. Papua New Guinea: Morobe, Mount Mis- 
im ( = Missim), 7,000 ft (2,135 m). 16 Januaiy 1933. 
Collector. H. Stevens. Original number 8. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. Taylor et al. (1982: 193) first synony- 
mized .stevensi with niobe, under the genus Rattus. 

Stenomys verecundus mollis Rummler, 

1935 

Z. Saugetierkunde, 10: 116, 31 

December. 

Holotype. MCZ 29905. Skin and skull. Adult female. 
Locality. Papua New Guinea: Morobe, Mount Mis- 
im ( = Missim), 5,850 ft (1,784 m). 14 April 1933. 
Collector. H. Stevens. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Flan- 
neiy (1995a: 346). 



Genus THAMNOMYS Thomas, 1 907 

Thamnomys venustus kivuensis G. M. 

Allen and Loverldge, 1942 

Bull Mus. Comp. Zool, 89: 192, 

Februaiy. 

Holotype. MCZ 39151. Skin and sk-ull. Adult female. 
Locality. Belgian Congo ( = Democratic Republic 
of Congo): (Sud-Kivu), Lake Kivu, Idjwi Island, 
Upper Mulinga, 6,500 ft (1,983 m). 2 March 1939. 
Collector. A. Loveridge. 

Condition. Skin complete. Skull partial (separated 
between nasal and frontal). Right and left angular 
processes of mandible chipped. 
Tiipe Series. Holotype only. 

Subfamily NESOMYINAE Forsyth Major, 
1897 

Genus NESOMYS Peters, 1870 

Nesomys lambertoni G. Grandidier, 1930c 
Bull. Acad. Malgache, n. sen, 11: 95 (for 
1928). 

Holotype. MCZ 45941. Skull and postcranial skele- 
ton. Adult inale. 

Locality. Malagasy Republic ( = Madagascar): west 
coast, vicinity of Maintirano. 

Collector No collection data available. Received by 
G. Grandidier from Academic Malgache, 1928. 
Condition. Skull partial (left zygomatic arch miss- 
ing, right jugal missing, several small holes). Post- 
cranial skeleton complete (small foot bones pre- 
svHTiably preserved in skin). 

Type Series. 2 paratopes, MCZ 45933 and 45934, 
poorly prepared skins (■with partial skulls within), 
both male. 

Comments. The skin belonging to the holotype is 
in the MNHN, Paris (M. Carleton, personal com- 
munication). Considered a valid species by Nowak 
(1999: 1434). 

Subfamily OTOMYINAE Thomas, 1897 

Genus OTOMYS F. Cuvier, 1824 

Otomys anchietae lacustris G. M. Allen 

and Loveridge, 1933 

Bull. Mus. Comp. Zool., 75: 120, 

Februaiy. 

= Otomys lacustris G. M. Allen and 

Loveridge, 1933. See Dieterlen and Van 

der Straeten (1992: 385). 

Holotype. MCZ 26358. Skin and sk-ull. Adult female. 
Locality. Tanganyika Territoiy (=Tanzania): north 
end of Lake Nyasa, Ukinga Mountains, Madehani, 
7,000 ft (2,135 m). 21 Februaiy 1930. 



Type Specimens of Recent Mammals • Helgen and McFadden 121 



Collector. A. Loveridge. 

Condition. Skin complete. Skiill partial (left tym- 
panic bulla damaged). Mandible disarticulated and 
left coronoid process chipped. 

Type Series. 17 paratypes; MCZ 26326, 26344- 
26351, 26353-26357, 26359, 26654, 26658; all rep- 
resented by skin and skull. 

Comments. Included in O. anchietae by Musser 
and Carleton (1993: 680) but treated as a full spe- 
cies by Nowak (1999: 1439). 

Otomys barbouri Lawrence and Loveridge, 

1953 

BulL Mus. Comp. ZooL, 110: 63, June. 

Holotijpe. MCZ 31369. Skin and skull. Adult male. 
Locality. Uganda: Mount Elgon, Kaburomi, 
1°14'N, 34°31'E, 10,500 ft (3,203 m). 28 December 
1933. 

Collector. A. Loveridge. 
Condition. Skin and skull complete. 
Type Series. 9 paratypes; MCZ 31371-31372, 
31376, 31421-31425, 31438 (4 males, 4 females, 
and 1 unsexed individual). 

Comments. Considered a valid species by Dieter- 
len and Van der Straeten (1992: 385) and Nowak 
(1999: 1439) but included in O. anchietae by Mus- 
ser and Carleton (1993: 680). 

Otomys uzungwensis Lawrence and 

Loveridge, 1953 

Bull Mus. Comp. ZooL, 110: 61, June. 

= Otomys typus uzungwensis Lawrence 

and Loveridge, 1953. See Hanney (1965: 

626). 

Holotype. MCZ 26645. Sldn and skull. Adult female. 
Locality. Tanganyika Territory (=Tanzania): Iringa 
District, Uzungwa Mountains, Dadaga, 6,000 ft 
(1,830 m). 31 December 1929. 
Collector. A. Loveridge. 
Condition. Skin and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. O. t. uzungwensis was retained as a val- 
id subspecies bv Ansell (1978: 78). 

Subfamily SIGMODONTINAE Wagner, 
1843 

Genus ISTHMOMYS Hooper and IVIusser, 
1964 

Megadontomys flavidus Bangs, 1902b 
Bull Mus. Comp. ZooL, 39: 27, ApriL 
= lsthmomys flavidus (Bangs, 1902). See 
Hooper and Musser (1964: 12). 

Holotype. MCZ 10331. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 4,000 ft (1,220 m). 12 April 1901. 



Collector W. W. Brown, Jr. Original number 28. 

Condition. Skin complete, widi small bald spots. 

Skull complete. Mandible disarticulated. 

Type Series. There is a large series of paratypes in 

the MCZ. 

Comments. Type species oihthmomys Hooper and 

Musser, 1964. I. flavidus was considered a valid 

species by Musser and Carleton (1993: 706) and 

Nowak (1999: 1357). 

Genus NECTOMYS Peters, 1861 

Nectomys squamipes amazonicus 
Hershkovitz, 1944 

Misc. Pub., Mus. ZooL, Univ. Mich., 58: 
47, 4 Januaiy. 

Holotype. MCZ 30820. Skin and skull. Male. 

Locality. Brazil: (Para), Rio Tapajos, Tauary (=Tau- 
ari). 23 January 1934. 

Collector A. M. Olalla. Original number 7312. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Hershkovitz examined 28 specimens, 
in the MCZ, AMNH, and FMNH. 
Comments. Retained as a valid subspecies by Er- 
nest (1986: 1). 

Genus NEOTOMA Say and Ord, 1825 

Neotoma abbreviata Goldman, 1909 

Proc. BioL Soc. Washington, 22: 140, 25 

June. 

= Neotoma lepida abbreviata Goldman, 

1909. See Burt (1932: 182). 

Holotype. MCZ 12260. Skin and skull. Adult male. 
Locality. (Mexico): Lower ( = Baja) California, San 
Francisco Island. 22 February 1909. 
Collector. W. W. Brown, Jn Original number 22. 
Condition. Skin and skull complete. 
Type Series. 9 paratypes; MCZ 12256-12259, 
12261—12265; all represented by skin and skull, 5 
females and 4 males. MCZ 12265 is now in the 
USNM. 

Comments. N. I. abbreviata was retained as a valid 
subspecies by Hall (1981: 755). 

Neotoma bella Bangs, 1899] 

Proc. New England ZooL Club, 1: 66, 31 

July. 

= Neotoma lepida lepida Thomas, 1893. 

See Goldman (1932: 62). 

Holotype. B5308. Skin and skull. Adult male. 

Locality. (United States): California, Riverside 
County, Palm Springs. 12 April 1896. 
Collector. E. C. Thurber Original number 623. 
Condition. Skin and skull complete. Mandible dis- 
articulated and left airgular pr'ocess chipped. 



122 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2 



Type Series. Holotype only. 

Neotoma distincta Bangs, 1903a 

Proc. Biol. Soc. Washington, 16: 89, 25 

June. 

= Neotoma mexicana distincta Bangs, 

1903. See Hall (1955: 329). 

Holotype. B9819. Skin and skull. Adult male. 

Locality. Mexico: Vera Cruz, near Jalapa, Texolo. 8 

March 1899. 

Collector. S. N. Rhoads. Original number 295. 

Condition. Skin and skull complete. 

Type Series. 4 paratypes; B9818, sldn and skull, 

male, B9820, skin and skull, female; B9821, skin 

and skull, male (exchanged to USNM); B9822, skin 

and skull, male. 

Comments. N. m. distincta was retained as a valid 

subspecies by Hall (1981: 761) and Cornely and 

Baker (1986: 1). 

Neotoma floridana rubida Bangs, 1 898b 
Proc. Boston Soc. Nat. Hist., 28: 185, 15 
March. 

Holotype. B2872. Skin and skull. Adult male. 

Locality. (United States): Louisiana, Terrebonne 
Parish, Gibson. 4 April 1895. 
Collector F. L. Small. Original number 1751. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 2 specimens are mentioned by num- 
ber in the original description: B2871, sldn and 
skull, adult male, and B2873, skin and skull, adult 
female. 

Comments. Retained as a valid subspecies by Hall 
(1981: 749). 

Genus NYCTOMYS 6e Saussure, 1860 

Nyctomys nitellinus Bangs, 1 902b 
Bull. Mus. Comp. ZooL, 39: 30, April. 
= Nyctomys sumichirasti nitellinus Bangs, 
1902. See Goldman (1916: 155). 

Holotype. MCZ 10249. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 4,000 ft (1,220 m). 8 Februaiy 1901. 
Collector. W. W. Brown, Jr. Original number 119. 
Condition. Skin and skull complete. 
Type Series. 5 paratypes; MCZ 10245, skin and 
skull, adult male; MCZ 10246, skin and skull, adult 
male; MCZ 10247, adult female, skin and skull; 
MCZ 10248, skin and skull, adult female; MCZ 
10250, sldn and skull, juvenile female. 
Comments. N. s. nitellinus was retained as a valid 
subspecies by Hall (1981: 630). 



Genus OECOMYS Thomas, 1906 

Oecomys trabeatus G. M. Allen and 

Barbour, 1923 

Bull. Mus. Comp. ZooL, 65: 262, 

Februaiy. 

= Oecomys bicolor trabeatus G. M. Allen 

and Barbour, 1923. See comments. 

Holotype. MCZ 19837. Skin and skull. Male. 
Locality. Panama: Rio Jesusito. 10 April 1922. 
Collector. T. Barbour and W. S. Brooks. Original 
number 2027. 

Condition. Skin complete. Skull partial (right zy- 
gomatic arch broken). Mandible disarticulated. 
Type Sei-ies. Holotype only. 

Comments. Following Hershkovitz (1960: 533), O. 
b. trabeatus was retained as a valid subspecies by 
Hall (1981: 619) under the genus Oryzomys. The 
use of Oecomys as a generic name follows Musser 
and Carleton (1993: 715). 

Oryzomys flavicans illectus Bangs, 1898k 

Proc. Biol. Soc. Washington, 12: 164, 10 

August. 

= Oecomys flavicans illectus (Bangs, 

1898k). See comments. 

Holotype. B8101. Sldn and skull. Adult female. 
Locality. Colombia: Magdalena, Santa Marta 
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 24 
March 1898. 

Collector W. W. Bi^own, Jr. Original number 89. 
Condition. Skin complete. Skull partial (right jugal 
missing). 

Tt/pe Series. There is a small series of paratypes in 
the MCZ. 

Comments. The use of Oecomys as a generic name 
follows Musser and Carleton (1993: 715). 

Genus OLIGORYZOMYS Bangs, 1 900 

Oryzomys navus Bangs, 1899a 

Proc. Biol. Soc. Washington, 13: 9, 31 

januaiy. 

= Oligoryzomys fulvescens (Saussure, 

1860). See Carleton and Musser (1989: 

70). 

Holotype. B8107. Skin and skull. Adult male. 

Locality. Colombia: Magdalena, Santa Marta 
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26 
March 1898. 

Collector W. W. Browni, Jr. Original number 154. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 9 paratypes; B8223-B8231; all repre- 
sented by sldn and skull, except B8228 (skin only); 
2 females and 7 males. B8224 is now in the USNM, 



Type Specimens of Recent Mammals * Helpen and McFadden 



123 



and B8223, B8226, B8230, and B8231 are now in 
the AMNH. 

Coininents. Oryzoini/s navus is the t)'pe species of 
Oligoryzomijs Bangs, 1899. 

Oryzomys (Oligoryzomys) vegetus Bangs, 

1902b 

Bull. Mus. Comp. ZooL, 39: 35, April. 

= Oligoryzomys vegetus (Bangs, 1902). 

See Musser and Carleton (1993: 718). 

Holotype. MCZ 10298. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Volcan de Chiriqui, 
Boquete, 4,000 ft (1,220 m). 16 April 1901. 
Collector. W. W. Brown, Jr. Original number 304. 
Condition. Skin complete. Skull partial (left jugal 
missing). Mandible disarticulated. 
Type Series. 12 paratypes; MCZ 10295 (now in 
USNM), 10297, 10300-10306, 10308-10310, 2 fe- 
males and 10 males, all represented by skin and 
skull except for 10308 (skin only). 
Comments. Considered a valid species by Musser 
and Carleton (1993: 718) and Nowak (1999: 1368). 
Carleton and Musser (1995) provided distribution- 
al information and support of specific status for O. 
vegetus. 

Genus OR VZOM VS Baird, 1858 

Oryzomys devius Bangs, 1902b 

Bull. Mus. Comp. ZooL, 39: 34, April. 

Holotype. MCZ 10324. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Volcan de Chiriqui, 
Boquete ( = Bajo Boquete), 5,000 ft (1,525 m). 29 
January 1901. 

Collector. W. W. Brown, jr. Original number 73. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paraty^aes; MCZ 10325, skin and 
skull, adult female; MCZ 10326, skin and skull, 
adult female; MCZ 10340, sldn and skull, adult 
male. 

Comments. Considered a valid species by Musser 
and Carleton (1993: 722) and Nowak (1999: 1366). 

Oryzomys palustris coloratus Bangs, 

1898b 

Proc. Boston Soc. Nat. Hist., 28: 189, 15 

March. 

= Oryzomys palustris natator Chapman, 

1893. See Wolfe (1982: 1). 

Holotype. B4470. Skin and skull. Adult male. 

Localitij. (United States): Florida, Monroe County, 

Cape Sable. 17 April 1895. 

Collector. C. L. Brownell. Original number 148. 

Condition. Sldn and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 



Oryzomys rostratus carrorum Lawrence, 

1947b 

Proc. New England ZooL Club, 24: 101, 

29 May. 

Holotype. MCZ 41280. Skin, skull, and postcranial 
skeleton. Adult male. 

Locality. Mexico: Tamaulipas, Rio Soto la Marina, 
Rancho Santa Ana, 8 miles (12.9 km) southwest of 
Padilla. 21 December 1941. 

Collector. G. H. Pournelle, Third University of 
Florida Mexican Expedition. Original number 196. 
Condition. Skin and skull complete. Mandible dis- 
articulated. Postcranial skeleton partial (right fore- 
limb, hindlimbs, pelvis, ribs, vertebral column). 
Type Series. 2 paratypes; MCZ 41281, skin, skull, 
and postcranial skeleton, female; MCZ 41282, sldn, 
skull, and postcranial skeleton, male. 
Comments. Retained as Oryzomys melanotis car- 
rorum by Hall (1981: 614). Musser and Carleton 
(1993: 724) regarded O. rostratus as distinct from 
O. melanotis. 

Genus PEROMYSCUS G\oger, 1841 

Hesperomys gossypinus LeConte, 1853 

Proc. Acad. Nat. Sci. Philadelphia, 6: 

411, 25 October. 

= Peromyscus gossypinus gossypinus 

(LeConte, 1853). See Rhoads (1896b: 

189). 

Lectotype. MCZ 5275. Skin and skull. Adult male. 
Locality. (United States): Georgia, Liberty County, 
Riceboro, probably on the LeConte Plantation near 
Riceboro. 13 September 1847. 
Collector J. E. LeConte. Formerly USNM 546. 
Condition. Skin complete. Skull partial (basioccip- 
ital and tympanic bullae broken). Mandible disar- 
ticulated. 

Type Series. Poole and Schantz (1942: 320) listed 
potential "cotypes," meaning syntypes. Osgood 
(1909: 136) mentioned that a specimen, number 
752 in the collection of the Academy of Natural 
Sciences in Philadelphia, possibly possessed type 
status, but this specimen was not mentioned by 
Koopman (1976). 

Comments. In the original description of gossypi- 
nus, measurements were given for a single speci- 
men only; however, in addition to this specimen, 
LeConte made reference to the coloration of 
"younger indixdduals." The measureinents given 
correspond to MCZ 5275 rather than to any other 
of LeConte "s specimens still in the USNM (see G. 
M. Allen, 1931: 262; Poole and Schantz, 1942: 
320). In addition, MCZ 5275 was the only speci- 
men of Peromyscus gossypinus from Georgia men- 
tioned in Baird's Mammals of North America 
(1858: 469). Because LeConte did not specify a ho- 
lotype, MCZ 5275, which seems to be the speci- 



124 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



men discussed and described by measurement in 
the original publication and the only adult speci- 
men mentioned, is here designated as lectotype. P. 
gossijpinus was considered a valid species by Mus- 
ser and Carieton (1993: 730) and Nowak (1999: 
1361). 

Hesperomys leucopus arcticus Mearns, 

1890 

Bull. Amer. Mus. Nat. Hist., 2: 285, 21 

Februaiy. 

Name preoccupied by Hesperomys 

arcticus Coues, 1 877 

Peromyscus maniculatus borealis Mearns, 

1911 

Proc. Biol. Soc. Washington, 24: 102, 15 

May. (Replacement name for 

Hesperomtjs leucopus arcticus Mearns, 

1890) 

Holotype. MCZ 5555. Skin and skull. Adult male. 
Locality. Canada: Northwest Territories, Mac- 
Kenzie, Fort Simpson, 7 September 1859. 
Collector. R. Kennicott. Original number 157. For- 
merly USNM 4531. 

Conclition. Skin complete. Skull partial (left jugal 
and process of squamosal missing, right process of 
squamosal missing, basioccipital and palatine miss- 
ing). Mandible disarticulated and both angular pro- 
cess missing. 

Type Series. Holot)'pe only. 

Comments. P. m. borealis was retained as a valid 
subspecies by Hall (1981: 672). 

Hesperomys sonoriensis nebrascensis 

Coues, 1877 

In Coues and J. A. Allen, Monographs N. 

Amer. Rodentia, U.S. Geol. Geogr. 

Survey Terr. Rep. Washington, 11: 79, 

August. 

= Peromyscus maniculatus nebrascensis 

(Coues, 1877). See Osgood (1909: 75). 

Sijntype. MCZ 5528. Skin and sk-ull. 

Locality. Nebraska (now Wyoming): (Converse 
County), Deer Creek. 19 Januaiy 1860. 
Collector F. V. Hayden. Original no. 80. Formerly 
USNM 4310. 

Condition. Sldn complete. Skull partial (posterior 
missing from parietal to palatine; both jugal and 
process of squamosal broken). Left angular process 
of mandible chipped, right side broken at cheek 
teeth. 

Type Series. 1 other syntype, present location un- 
known, formerly in die USNM. 
Comments. Hesperomys sonoriensis nebrascensis 



Baird, 1858 is a nomen nudum. Osgood (1909: 78) 
and Mearns (1911: 102) designated Deer Creek, 
Nebraska, as the type locaHty. Jones (1958: 107- 
111) reviewed the systematic histoiy oi nebrascen- 
sis, and Jones and Mursaloglu (1961: 101-103) re- 
ported the discovery of one of the two original syn- 
types in the MCZ. The location of the other syn- 
tN-pe, if it still exists, is unknown. 

Peromyscus anastasae Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 195, 15 

March. 

= Peromyscus gossypinus anastasae 

Bangs, 1898. See Osgood (1909: 141). 

Holotype. B7179. Skin and skull. Adult female. 
Locality. (United States): Florida, St. Johns Coun- 
ty, Anastasia Island, Point Romo. 15 February 
1897. 

Collector O. Bangs. Original number 2. 
Condition. Sldn complete, skull partial (premaxilla 
to frontal in fragments). Right mandible missing, 
left present. 

Type Series. There is a small series of paratypes in 
the MCZ. 

Comments. P. g. anastasae was retained as a valid 
subspecies by Hall (1981: 689). 

Peromyscus bellus Bangs, 1896h 

Proc. Biol. Soc. Washington, 10: 137, 28 

December. 

= Peromyscus attwateri ^. A. Allen, 1895. 

See Schmidly (1973: 125). 

Holott/pe. B5483. Skin and skull. Adult male. 

Locality. (United States): Indian Territory 
( = Oklahoma), (Adair County), Stilwell. 15 August 
1896. 

Collector T Surber. Original number 67. 
Condition. Skin and sk-ull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B5484, skin and skull, 
adult female. 

Peromyscus cacabatus Bangs, 1 902b 
Bull. Mus. Comp. Zool, 39: 29, April. 
= Peromyscus mexicanus (Saussure, 
1860). See Huckaby (1980: 15). 

Holotype. MCZ 10225. Sldn and skull. Adult female. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 5,000 ft (1,525 m). 22 April 1901. 
Collector. W. W. Brown, Jr. Original number 331. 
Condition. Skin complete. Skull partial (left jugal 
missing). Mandible disarticulated. 
Type Series. There is a very large series of paraty- 
pes in the MCZ. 



Type Specimens of Recent Mammals • Helgen and McFadden 125 



Peromyscus canadensis abietorum Bangs, 

1896d 

Proc. Biol. Soc. Washington, 10: 49, 9 

March. 

= Peromyscus maniculatus abietorum 

Bangs, 1896. See Osgood (1909: 45). 

Holotype. B2205. Skin and sknll. Adult female. 

Locality. (Canada): Nova Scotia, James River. 8 

August 1894. 

Collector. C. H. Goldthw^aite. 

Condition. Sldn complete. Skull partial (right jugal 

missing). 

Type Series. There is a series of paratypes in the 

MCZ. 

Comments. P. m. abietorum was retained as a valid 

subspecies by Hall (1981: 670). 

Peromyscus canadensis argentatus 

Copeland and Church, 1906 

Proc. Biol. Soc. Washington, 19: 122, 6 

September. 

= Peromyscus maniculatus argentatus 

Copeland and Church, 1906. See Osgood 

(1909: 46). 

Holotype. MCZ 54627. Skin and skull. Adult male. 
Locality. (Canada): New Brunswick, Grand Manan 
Island, Grand Harbor. 19 September 1905. 
Collector M. L. Church and M. Copeland. Original 
number 168. 

Condition. Skin and skull complete. 
Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. P. m. argentatus was retained as a valid 
subspecies by Hall (1981: 671). 

Peromyscus crinitus scitulus Bangs, 1899j 

Proc. New England Zool. Club, 1: 67, 31 

July. 

= Peromyscus crinitus crinitus (Merriam, 

1891). See Osgood (1909: 229). 

Holotype. B9175. Skin and skull. Adult male. 

Locality. (United States): Nevada, Douglas County, 

Gardnerville. 13 July 1898. 

Collector W. W. Price and R O. Simons. Original 

number 1571. 

Condition. Skin complete. Skull partial (right and 

left jugal missing). 

Type Series. 7 paratypes; B9177-B9180, B9183- 

B9185; all represented by skin and skull, 4 females 

and 3 males. 



Peromyscus gossypinus nigriculus Bangs, 

1896e 

Proc. Biol. Soc. Washington, 10: 124, 5 

November. 

= Peromyscus gossypinus gossypinus 

(LeConte, 1853). See Osgood (1909: 136). 

Holotype. B2731. Sldn and skull. Adult female. 

Locality. (United States): Louisiana, Plaquemines 
Parish, Burbridge. 30 Januaiy 1895. 
Collector F. L. Small. Original number 1547/156. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a veiy large series of paraty- 
pes in the MCZ. 

Peromyscus gossypinus palmarius Bangs, 

1896e 

Proc. Biol. Soc. Washington, 10: 124, 5 

November. 

Holotype. B3224. Skin and skull. Adult female. 
Locality. (United States): Florida, Brevard County, 
Oak Lodge on East Peninsula opposite Micco. 23 
Februaiy 1895. 
Collector O. Bangs. 
Condition. Skin and skull conaplete. 
Type Series. There is a very large series of paraty- 
pes in the MCZ. 

Comments. Retained as a valid subspecies by Hall 
(1981: 690). 

Peromyscus insulanus Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 196, 15 

March. 

= Peromyscus gossypinus anastasae 

Bangs, 1898. See Osgood (1909: 141). 

Holotype. B6438. Skin and skull. Adult male. 

Locality. (United States): Georgia, Camden Coun- 
ty, Cumberland Island, north end. 10 April 1897. 
Collector W. W. Brown, Jr. Original number 804. 
Condition. Skin complete. Skull partial (right jugal 
missing) . 

Type Series. There is a large series of paratypes in 
the MCZ. 

Peromyscus leucopus ammodytes Bangs, 

1905a 

Proc. New England Zool. Club, 4: 14, 28 

Februaiy. 

Holotype. B828. Skin and skull. Adult male. 

Locality. (United States): Massachusetts, Barnsta- 
ble County, Monomoy Island. 28 December 1893. 
Collector G. S. Miller, Jr. and O. Bangs. Original 
number 8. 
Condition. Skin and skull complete. 



126 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Type Series. There is a series of paratypes in the 

MCZ. 

Comments. Retained as a vaUd subspecies by Hall 

(1981: 685). 



Peromyscus leucopus fusus Bangs, 1 905a 
Proc. New England Zool. Club, 4: 13, 28 



Februaiy. 



Holotijpe. B9737. Skin and skull. Adult male. 

Localitij. (United States): Massachusetts, Dukes 

County, Martha's Vineyard, W. Tisbury. 17 June 

1899. 

Collector. O. Bangs. Original number 1. 

Condition. Skin and skull complete. 

Type Series. There is a small series of paratypes in 

the MCZ. 

Comments. Retained as a valid subspecies by Hall 

(19S1: 686). 



Peromyscus oreas Bangs, 1 898f 

Proc. Biol. Soc. Washington, 12: 84, 24 

March. 



Holotype. B3696. Skin and skull. Adult female. 

Locality. Canada: British Columbia, near boundaiy 

of Whatcom County, Washington, 49th parallel. 

Mount Baker Range, 6,500 ft (1,983 m). 29 August 

1896. 

Collector A. C. Brooks. Original number 745. 

Condition. Sldn and skull complete. 

Type Series. There is a large series of paratypes in 

the MCZ. 

Comments. Considered a valid species by Musser 

and Carleton (1993: 734) and Nowak (1999; 1361). 

Peromyscus phasma Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 199, 15 

March. 

= Peromyscus poloniotus phasma Bangs, 

1898. See Osgood (1909: 107). 

Holotype. B7175. Skin and skull. Adult female. 

Locality. (United States): Florida, St. Johns Coun- 
ty, Anastasia Island, Point Romo. 14 Februaiy 
1897. 

Collector O. Bangs. Original number 3. 
Condition. Skin complete. Skull partial (parietal, 
tympantic bulla, left jugal, and process of squa- 
mosal broken). Mandible disarticulated. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. P. p. phasma was retained as a valid 
subspecies by Hall (1981: 668). 



Peromyscus subgriseus arenarius Bangs, 

1898b 

Proc. Boston Soc. Nat. Hist., 28: 202, 

March. 

Name preoccupied by Peromyscus 

eremicus arenarius Mearns, 1 896. 

Peromyscus subgriseus baliolus Bangs, 

1898n 

Science, n. sen, 8: 215, 19 August. 

(Replacement name for Peromyscus 

subgriseus arenarius Bangs, 1898) 

= Peromyscus polionotus polionotus 

(Wagner, 1843). See Osgood (1909: 104). 

Holotype. B5925. Skin and skull. Adult male. 

Locality. (United States): Georgia, Scriven County, 

Hursman's Lake (Savannah River), near Bascom. 

15 December 1896. 

Collector W. W. Brown, Jr Original number 60. 

Condition. Skin and skull complete. 

Type Series. There is a large series of paratypes in 

the MCZ. 

Peromyscus subgriseus rhoadsi Bangs, 

1898b 

Proc. Boston Soc. Nat. Hist., 28: 201, 15 

March. 

= Peromyscus polionotus riioadsi Bangs, 

1898. See Osgood (1909: 107). 

Holotype. B6980. Skin and skull. Adult male. 

Locality. (United States): Florida, Hillsborough 

County, head of Anclote River. 23 May 1895. 

Collector. W. S. Dickinson. 

Condition. Sldn and skvdl complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Comments. P. p. rhoadsi was retained as a valid 

subspecies by Hall (1981: 669). 

Peromyscus texanus saturatus Bangs, 

1897a 

Amer. Nat. 31: 75, 1 Januaiy. 

= Peromyscus maniculatus saturatus 

Bangs, 1897. See Osgood (1909: 61). 

Holotype. B2581. Skin and skull. Adult male. 

Locality. Canada: British Columbia, Saturna Is- 
land. 31 Januaiy 1894. 

Collector W. C. Colt. Original number 128. 
Condition. Skin and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. P. m. saturatus was retained as a valid 
subspecies by Hall (1981: 682). 



Type Specimens of Recent Mammals • Helpen and McFadden 



127 



Genus REITHRODONTOMYS Giglioli, 
1873 

Reithrodontomys australis vulcanius 
Bangs, 1902b 

Bull. Mus. Comp. Zool, 39: 38, April. 
= Reithrodontomys sumichrasti vulcanius 
Bangs, 1902. See Hooper (1952: 83). 

Holotijpe. MCZ 10281. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Volcan de Chiriqui, 
10,300 ft (3,142 m). 26 May 1901. 
Collector. W. W. Brown, Jr. Original number 500. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. R. s. vulcanius was retained as a valid 
subspecies by Hall (1981: 644). 

Reittirodontomys creper Bangs, 1902b 
Bull. Mus. Comp. Zool, 39: 39, April. 

Holotype. MCZ 10284. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Volcan de Chiriqui, 
11,000 ft (3,355 m). 2 June 1901. 
Collector. W. W. Brown. Original number 504. 
Condition. Skin and skull complete. Mandible dis- 
articulated. Left lower incisor missing. 
Type Series. Holotype only. 

Comments. Considered a valid species by Musser 
and Carleton (1993: 740) and Nowak (1999: 1364). 

Reithrodontomys lecontii impiger Bangs, 

1898m 

Proc. Biol. Soc. Washington, 12: 167, 10 

August. 

= Reithrodontomys humulis humulis 

(Audubon and Bachman, 1841). See Hall 

and Kelson (1959: 584). 

Holotype. B7784. Skin and skull. Adult male. 

Locality. (United States): West Virginia, Green- 
briar County, White Sulphur Springs, 2,000 ft (610 
m). 27 Febmaiy 1898. 
Collector. T. Surber. Original number 466. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 2 paratypes; B6931 (not B6932, as 
stated in the original description), skin and skull, 
adult male; B7785, skin and skull, adult female. 

Genus SCOT/A/OM/S Thomas, 1913 

AlKodon teguina apricus Bangs, 1 902b 
Bull. Mus. Comp. Zool, 39: 40, April. 
= Scotinomys teguina apricus (Bangs, 
1902). See Thomas (1913: 409). 

Holotype. MCZ 10236. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Boquete ( = Baio Bo- 
quete), 4,000 ft (1,220 m). 24 February 1901. 



Collector. W W. Brown, Jr. Original number 192. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 4 paratypes; MCZ 10234, skin and 
skull, adult male; MCZ 10235, skin, adult female; 
MCZ 10237, skin and skull, adult female; MCZ 
10238, sldn and skull, adult female. 
Comments. S. t. apricus was retained as a valid sub- 
species by Hooper (1972: 21) and Hall (1981: 734). 

AI<odon xerampelinus Bangs, 1 902b 
Bull. Mus. Comp. Zool., 39: 41, April. 
= Scotionomys xerampelinus (Bangs, 
1902). See Thomas (1913: 409). 

Holotype. MCZ 10240. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Volcan de Chiriqui, 
10,300 ft (3,142 m). 26 May 1901. 
Collector W W Brown, Jr. Original number 501. 
Condition. Sldn complete. Skull partial (right jugal 
missing). Mandible disarticulated. 
Type Series. 2 paratypes; MCZ 10239, skin and 
skull, adult male; MCZ 10241, skin and skull, adult 
male. 

Comments. S. xerampelinus was considered a valid 
species by Musser and Carleton (1993: 746) and 
Nowak (1999: 1355). 

Genus SIGMODON Say and Ord, 1825 

Sigmodon austerulus Bangs, 1 902b 
Bull. Mus. Comp. Zool, 39: 32, April. 
= Sigmodon hispidus borucae J. A. Allen, 
1897. See Hall and Kelson (1959: 672). 

Holotype. MCZ 10288. Sldn and skull. Adult male. 
Locality. Panama: Chiiiqui, Volcan de Chiriqui, 
10,000 ft (3,050 m). 1 June 1901. 
Collector W W Brown, Jr. Original number 503. 
Condition. Skin complete. Skull partial — according 
to original description, "unfortunately it [the skull] 
was broken by the trap directly across the orbits." 
Tijpe Series. Holotype only. 

Comments. For comments regarding the type lo- 
cality, see Enders (1953a: 508-509), who suggested 
that a native helper of W. W Brown, Jr, transplant- 
ed a specimen of S. [h.] borucae, trapped during a 
trip to his home in the lowlands near Boruca, to 
the top of the Volcan, "after Brown had promised 
a bottle of wine to any man who captured a Sig- 
modon at that altitude." 

Sigmodon hispidus exsputus G. M. Allen, 

1920b 

J. Mammal., 1: 236, 4 December. 

Holotype. MCZ 18100. Skin and skull. Adult male. 
Locality. (United States): Florida, Monroe County, 
Big Pine Key 16 April 1920. 
Collector W S. Brooks. Original number 1936. 



128 Bidletm Museum of Comparative Zoology, Vol. 157, No. 2 



Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paraty][De; MCZ 18101, skin and 
skull, adult male. 

Comments. Retained as a valid subspecies by Hall 
(1981: 737). 



Sigmodon hispidus furvus Bangs, 1 903b 
Bull. Mus. Comp. Zool., 39: 158, July. 



Holottjpe. MCZ 10665. Skin and skull. Male. 

Locality. Honduras: Atlantida, La Ceiba. 16 Janu- 
ary 1902. 

Collector. W. W. Brown, Ji". Original number 4. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Hall 
(1981: 738). Sigmodon hispidus fei'vidus Lydekker, 
1904 (p. 34) was a misspelling and thus accidental 
renaming of Sigmodon hispidus furvtis Bangs, 
1903. 



Sigmodon hispidus spadicipygus Bangs, 

1898b 

Proc. Boston Soc. Nat. Hist., 28: 192, 15 

March. 



Holotype. B4477. Skin and skull. Adult female. 

Locality. (United States): Florida, Monroe County, 
Cape Sable. 18 April 1895. 

Collector C. L. Brownell. Original number 153. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a series of paratypes in the 
MCZ. 

Comments. Retained as a valid subspecies by Hall 
(1981: 739). 



Sigmodon sanctae-martae Bangs, 1898q 

Proc. Biol. Soc. Washington, 12: 189, 30 

December. 

= Sigmodon hispidus hirsutus (Burmeister, 

1854). See Cabrera (1961: 508). 



Holotype. B8105. Skin and skull. Adult male. 

Locality. Colombia: (Magdalena), (Santa Marta 
Mountains), Pueblo Viejo (not far from the source 
of Rio Ancho), 8,000 ft (2,440 m). 23 March 1898. 
Collector. W. W. Brown, Jr. Original number 73. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B8250, skin and skull, 
adult male. 



Genus THOIVI ASOIVIYS Coues, 1884 

Oryzomys (Erioryzomys) monochromos 

Bangs, 1900b 

Proc. New England Zool. Club, 1: 97, 23 

February. 

= Thomasomys monochromos (Bangs, 

1900). See Gardner and Patton (1976: 

26). 

Holotype. B8348. Sldn and skull. Adult male. 

Locality. Colombia: Magdalena, Santa Marta 
Mountains, Paramo de Macotania, 11,000 ft (3,355 
m). 7 March 1899. 

Collector W. W. Brown, Jr. Original number 93. 
Condition. Skin complete. Skull partial (right jugal 
missing). Majidible disarticulated. 
Type Series. 4 paratypes; B8248, juvenile male, 
B8345, adult female, B8346, adult female, B8347, 
adult male; all represented by sldn and skull. 
Comments. Considered a valid species by Musser 
and Carleton (1993: 750) and Nowak (1999: 1362). 
Paramo de Macotama is in La Guajira (Paynter 
1997: 261). 

Genus ZYGODONTOMYS ^. A. Allen, 
1897 

Zygodontomys seorsus Bangs, 1901b 
Amer. Nat., 35: 642, 22 August. 
= Zygodontomys brevicauda cherriei (J. A. 
Allen, 1895). See Voss (1991: 59). 

Holotype. B8490. Skin and skull. Adult male. 

Locality. Panama: Gulf of Panama, San Miguel Is- 
land, Isla del Rey 5 May 1900. 
Collector W. W. Brown, Jr Original number 147. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. Bangs based the description on a se- 
ries of 68 specimens, most of which are still in the 
MCZ. 

Family PEDETIDAE Gray, 1825 

Genus PEDETES llliger, 1811 

Pedetes cafer taborae G. M. Allen and 

Loveridge, 1927 

Proc. Boston Soc. Nat. Hist., 38: 438, 23 

December. 

= Pedetes capensis (Forster, 1778). See 

Misonne (1974: 8). 

Holotype. MCZ 23080. Skin and skull. Adult female. 
Locality. Tanganyika Territory (=Tanzania): Ta- 
bora, Unyamwezi, 4,000 ft (1,220 m). 18 November 
1921. 
Collector A. Loveridge. Original number R6915. 



Type Specimens of Recent Mammals • Helgen and McFadden 129 



Condition. Sldn complete, with small bald spots on 
dorsum. Skull complete. 
Type Series. Holotype only. 

Family MYOXIDAE Gray, 1821 

Genus GRAPHIURUS Smuts, 1832 

Aethoglis hueti argenteus G. M. Allen, 

1936 

J. Mammal, 17: 293, 14 August. 

= Graphiurus hueti argenteus (G. M. Allen, 

1936). See Hayman and Holt (1940: 

608). 

Holotype. MCZ 17920. Skin and skull. Adult female. 
Locality. Cameroons ( = Cameroon): (Kribi), Lolo- 
dorf. 17 March 1911. 

Collector. G. Schwab. Original number 1. 
Condition. Sldn and skull complete. 
Type Series. 2 paratypes; MCZ 17607, sldn and 
skull, adult female; and a specimen in the USNM, 
number 125434. 

Comments. Rosevear considered G. h. argenteus 
"possibly valid as a race" (1969: 501). 

Claviglis soleatus collaris G. M. Allen and 

Loveridge, 1933 

Bull. Mus. Comp. Zool., 75: 122, 

Februan'. 

= Graphiurus lorraineus Dollman, 1910. 

See Holden (1993: 764). 

Holotype. MCZ 26373. Skin and skull. Adult female. 
Locality. Tanganyika Territory (=Tanzania): (Irin- 
ga), north end Lake Nyasa, Uldnga Mountains, Ma- 
dehani, 7,000 ft (2,135 m). 24 Februaiy 1930. 
Collector. A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 2 paratypes; MCZ 26374, skin and 
skull, male; MCZ 26375, skin and skull, female. 

Graphiurus microtis griseus G. M. Allen, 

1912a 

Bull. Mus. Comp. Zool, 54: 440, April. 

= Grapfiiurus murinus griseus G. M. Allen, 

1912. See Hayman and Holt (1940: 610). 

Holotype. MCZ 8244. Skin and skull. Adult male. 
Locality. British East Africa ( = Kenya): northern 
Guaso Nyiro River ( = Ewaso Ngiro), 4,000 ft (1,220 
m). 25 July 1909. 

Collector G. M. Allen. Original number 51. 
Condition. Skin and skull complete. 
Type Series. 2 paratypes; MCZ 8248, skin and skull, 
male; MCZ 8249, skin and skaiU, adult female. 



Grapiiiurus scfiwabi G. M. Allen, 1912a 
Bull. Mus. Comp. Zool., 54: 441, April. 
= Grapiiiurus surdus Dollman, 1912. See 
Holden (1993: 765). 



Holotype. MCZ 8607. Skin and skull. Juvenile. 
Locality. Cameroun ( = Cameroon), Kribi. 1911. 
Collector G. Schwab. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Family BATHYERGIDAE Waterhouse, 
1841 



Genus CRYPTOIVIYS Gmy, 1864 

Cryptomys tiottentotus occlusus G. M. 

Allen and Loveridge, 1933 

Bull. Mus. Comp. Zoo)., 75: 125, 

Februaiy. 

= Cryptomys tiottentotus wfiytei (Thomas, 

1897). See Honeycutt et al. (1991: 51). 

Holotype. MCZ 26557. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (=Tanzania): (Irin- 
ga), Ugungwe Mountains, Kigogo, 6,000 ft (1,830 
m). 18 January 1930. 
Collector. A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 15 paratypes; MCZ 26558-26572, all 
represented by skin and skull. 

Genus HETEROCEPHALUS Ruppell, 
1842 

l-ieterocepiiaius stygius G. M. Allen, 

1912a 

Bull. Mus. Comp. Zool, 54: 444, April. 

= l-leterocepiiaius giaber Ruppell, 1842. 

See HoUister (1919: 160). 



Holotype. MCZ 12470. Body in alcohol, skull e.xtract- 
ed. Adult female. 

Locality. British East Africa ( = Kenya): northern 
Guaso Nyiro ( = Ewaso Ngiro), Neumann's Boma. 6 
August 1909. 
Collector G. M. Allen. 

Condition. Alcoholic. Skull complete. Mandible 
disarticulated. 
Type Series. Holotype only. 



130 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Family ERITHIZONTIDAE Bonaparte, 
1845 

Genus ERITHIZONF. Cuvier, 1822 

Erethizon dorsatus picinus Bangs, 1900c 

Proc. New England Zool. Club, 2: 37, 20 

September. 

= Erithizon dorsatum dorsatum (Linnaeus, 

1758). See Hai-per (1961: 90). 

Holotype. B8839. Sldn and skull. Adult male. 

Locality. (Canada): Labrador, Strait of Belle Isle, 
Lance ( = Lanse) au Loup. 16 Februaiy 1899. 
Collector. E. Doane. 
Condition. Skin and skull complete. 
Type Series. 1.5 paratypes; B8832-B8838, B8840- 
B8847; all represented by skin and skull, 8 females 
and 7 males. Bangs had also a skull of picimis 
(B8831) from Black Bay, Labrador, at the time of 
description, not mentioned in the original publi- 
cation. 

Family DASYPROCTIDAE Gray, 1825 

Genus DASYPROCTA llliger, 1811 

Dasyprocta callida Bangs, 1 901 b 
Amer. Nat., 35: 635, 22 August. 
= Dasyprocta punctata callida Bangs, 
1901. See Kellogg (1946: 59). 

Holotype. B8443. Skin and skull. Adult male. 

Locality. Panama: (Panama), San Miguel Island. 8 

May 1900. 

Collector. W. W. Brown, Jr Original number 171. 

Condition. Skin and skull complete. 

Type Series. 5 paratypes; B8442, B8444-B8447; all 

represented by skin and skull, 3 females and 2 

males. 

Comments. D. p. callida was retained as a valid 

subspecies by Hall (1981: 860). 

Dasyprocta colombiana Bangs, 1 898k 

Proc. Biol. Soc. Washington, 12: 163, 10 

August. 

= Dasyprocta punctata colombiana (Gray, 

1898). See Cabrera (1961: 589). 

Holotype. B8008. Skin and skull. Adult female. 

Locality. Colombia: Magdalena, Santa Marta. 6 
January 1898. 

Collector. W. W. Brown, Jr. Original number 37. 
Condition. Sldn and skull complete. 
Type Series. 1 paratype; B8113, skin and skull, ju- 
venile male. 

Comments. D. p. colombiana was retained as a val- 
id subspecies by Cabrera (1961: .589). 



Dasyprocta noblei G. M. Allen, 1914e 

Proc. New England Zool. Club, 5: 69, 7 

October. 

= Dasyprocta leporina noblei G. M. Allen, 

1914. See Woods (1993: 781). 

Holotype. MCZ 15936. Sldn and skull (and atias). 
Adult female. 

Locality. Guadeloupe Island: Goyave. 22 August 
1914. 

Collector G. K. Noble. 

Condition. Sldn complete. Skull partial (bone miss- 
ing from nasal and frontal). 

Type Series. 1 paratype; MCZ 15937, sldn and 
skull, subadult female. 

Dasyprocta punctata nuchalis Goldman, 

1917 

Proc. Biol. Soc. Washington, 30: 113, 23 

May. 

Holotype. MCZ 1008 L Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Divala. 30 November 
1900. 

Collector W. W. Brown, Jr. Original number 17. 
Condition. Skin and skull complete. 
Type Series. 4 paratypes; MCZ 10080, skin and 
skull, adult male; 10084, skull; 10175, skin and 
skull, adult female; 10176, sldn and skull, adult 
male. 

Comments. Retained as a valid subspecies by Hall 
(1981: 860). 

Family AGOUTIDAE Gray, 1821 

Genus AGOUTI Lacepede, 1799 

Agouti paca virgatus Bangs, 1902b 
Bull. Mus. Comp. Zool., 39: 47, April. 

Holotype. MCZ 10079. Sldn and skull. Adult male. 
Locality. Panama: Chiriqui, Divala. 16 December 
1900. 

Collector W. W. Brown, Jr Original number 21. 
Condition. Sldn and skull complete. 
Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Hall 
(1981: 8.58) and Perez (1992: 1). 

Family OCTODONTIDAE Waterhouse, 

1839 

Genus TYMPANOCTOMYS Yepes, 1941 

Octomys barrerae Lawrence, 1941 
Proc. New England Zool. Club, 18: 43, 
28 Januaiy. 

= tympanoctomys barrerae (Lawrence, 
1941). See Yepes (1942: 75). 

Holotype. MCZ 39716. Skin and skull. Aduk male. 
Locality. Argentina: Mendoza Province, La Paz. 
April 1939. 



Type Specimens of Recent Mammals • Helpen and McFadden 



131 



Collector. J. M. de la Barrera. Original number 

A31. 

Condition. Sldn and skull complete. 

Type Series. Holotype only. 

Comments. Type species of Tyinpanoctomys Yepes, 

1941. T. barrerae was considerd a valid species by 

Woods (1993: 789) and Nowak (1999: 1683). 

Family ECHIMYIDAE Gray, 1825 

Genus BOROMYS M\\\er, 1916 

Boromys torrei G. M. Allen, 1917a 
Bull. Mus. Comp. Zool., 61: 6, Januaiy. 

Holotype. VP 9601. Palate. 

Locality. Cuba: Matanzas Province, cave in Sierra 
de Hato Nuevo. 
Collector C. de la Torre. 

Condition. Condition as described in original — 
"palate with root of right z}'gomatic arch, pni^ and 
alveolar row of right side, i)i' and posterior part of 
alveolar row of left side." 

Type Series. Paratype material consists of 8 lower 
jaws and 2 separate lower molars; in the MCZ. 
Comments. Considered a valid species by Woods 
(1993: 799) and Nowak (1999: 1703) but almost 
certainly extinct. This specimen is stored in the 
Vertebrate Paleontology Department of tlie MCZ. 

Genus DIPLOMYS Thomas, 1916 

Loncheres labilis Bangs, 1901b 
Amer. Nat., 35: 638, 22 August. 
= Diplomys labilis (Bangs, 1901). See 
Thomas (1916: 296). 

Holotype. B8480. Skin and skull. Adult male. 

Locality. Panama: (Panama), Gulf of Panama, San 
Miguel Island. 26 April 1900. 

Collector. W W Brown, Jr. Original ninuber 103. 
Condition. Skin and skull complete. 
Type Series. There is a small series of paratypes in 
the MCZ and FMNH. 

Comments. D. labilis was considered a valid spe- 
cies by Woods (1993: 791) and Nowak (1999: 
1695). 

Genus PROECHIMYS ^. A. Allen, 1899 

Proechimys burrus Bangs, 1901b 
Amer. Nat., 35: 640, 22 August. 
= Proechimys semispinosus burrus Bangs, 
1901. See Goldman (1920: 120). 

Holotype. B8458. Skin and skull. Adult male. 

Locality. Panama: (Panama), Gulf of Panama, San 
Miguel Island. 30 April 1900. 
Collector WW. Bro\\ai, Jr. Original number 130. 
Condition. Skin and skull complete. Bight dentary 
broken at pnij. 



Type Series. There is a large series of paratypes in 
the MCZ. 

Comments. P. s. burnis was retained as a valid sub- 
species bv Hall (1981: 873). 

Proechimys gorgonae Bangs, 1905b 
Bull. Mus. Comp. Zool., 46: 89, June. 

Holotype. MCZ 10828. Sldn and skull. Adult male. 
Locality. Colombia: (Cauca), Gorgona Island. 2 
July 1904. 

Collector W. W Brown, Jr. Original number 25. 
Condition. Skin and skull complete. 
Tijpe Series. 6 paratypes; MCZ 10829-10834, all 
represented by skin and skull, 1 female and 5 
males. MCZ 10830 and 10834 are now in the 
FMNH. 

Comments. Treated as a separate species by Woods 
(1993: 796) but included in P. cayennensis by No- 
wak (1999: 1689). 

Proechimys guyannensis hyleae Moojen, 

1948 

Univ. Kansas Publ. Mus. Nat. Hist., 1: 

361, 10 December. 

= Proechimys cayennensis hyleae Moojen, 

1948. See comments. 

Holotype. MCZ 30887. Skin and skull. Adult male. 
Locality. Brazil: Para, Porto de Moz, Rio Tapajoz 
(=Tapajos), Tauari (=Tauaiy), 87 km south of San- 
tarem. 19 January 1934. 

Collector A. M. Olalla. Original number 7288. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 20 paratypes (19 at MCZ and 1 at 
FMNH), represented by sldn and skull. 
Comments. Retained as a valid subspecies by Ca- 
brera (1961: 521). The use of the specific name 
cayennensis over guyannensis follows Woods 
(1993: 795). 

Family CAPROMYIDAE Smith, 1842 

Genus CAPROMYS Desmarest, 1822 

Capromys pilorides relictus G. M. Allen, 

1911a 

Bull. Mus. Comp. Zool, 54: 207, July. 

Holotype. MCZ 10996. Skin and skull. Adult male. 
Locality. Cuba: Isle of Pines ( = Isla de Pinos), Nue- 
va Gerona, Casas Mountains. 10 March 1902 
Collector. W. R. Zappey. 

Condition. Sldn complete. Skull partial (bone miss- 
ing from palatine to occipital). 

Type Series. 1 paratype; MCZ 10997, sldn and 
skull. 

Comments. Retained as a valid subspecies by Hall 
(1981: 863). 



132 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Genus GEOCAPROMYS Chapman, 1901 

Geocapromys cubanus G. M. Allen, 1917a 
Bull. Mus. Comp. Zool. 61: 9, Januaiy. 
= Geocapromys columbianus (Chapman, 
1892). See G. M. Allen (1918b: 145). 

Holotype. VP 9602. Part of right lower ramus. Juve- 
nile. 

Locality. Cuba: Matanzas Province, cave in Sierra 
de Hato Neuvo. 
Collector. C. de la Torre. 

Condition. Portion of ramus includes incisor and 
three anterior cheekteeth. 

Type Series. Allen mentions in the original descrip- 
tion, "Five palates with teeth, about 15 jaw frag- 
ments mostly with teeth, and numerous other frag- 
ments." This material is in the MCZ. 
Comments. The type material o{ cubanus is stored 
in the Vertebrate Paleontology Department of the 
MCZ. 

Geocapromys ingrahami abaconis 
Lawrence, 1934 

Occas. Pap. Boston See. Nat. Hist., 8: 
190, 7 November. 

Holotype. VP 2108. Left lower ramus. Adult. 

Locality. Bahamas: Great Abaco Island, Hole in 
the Wall, Imperial Lighthouse Caves. 1934. 
Collector F. Rainey. 
Condition. Ramus intact. 

Type Series. The parats'pe series is a collection of 
cranial and mandibular fragments, described in the 
original description and housed in the MCZ. 
Comments. G. i. abaconis was retained as a valid 
subspecies by Hall (1981: 866) under the genus 
Capromys. Woods (1993: 800) employed the genus 
Geocapromys. G. i. abaconis is extinct. The type 
material of abaconis is stored in the Vertebrate Pa- 
leontology Department of the MCZ. 

Geocapromys ingrahami irrectus 
Lawrence, 1934 

Occas. Pap. Boston Soc. Nat. Hist., 8: 
190, 7 November. 

Holotype. VP 2107. Right lower ramus. Adult. 

Locality. Bahamas: Crooked Island, Gordon Hill 
Caves, "Burial Cave No. 1." 1934. 
Collector. F. Rainey. 
Condition. Ramus intact. 

Tijpe Series. The paratype series is a collection of 
cranial and mandibular fragments, described in the 
original description and housed in the MCZ. 
Comments. G. i. irrectus was retained as a valid 
subspecies by Hall (1981: 866) vmder the genus 
Capromys. Woods (1993; 800) employed tlie genus 
Geocapromys. G. i. irrectus is extinct. The type ma- 



terial of irrectus is stored in the Vertebrate Pale- 
ontology Department of tlie MCZ. 

Genus MESOCAPROMYS Marona, 1970 

Capromys nana G. M. Allen, 1917b 

Proc. New England Zool. Club, 6: 54, 28 

March. 

= Mesocapromys nanus (G. M. Allen, 

1917). See Kratochvil et al. (1978: 15). 



Holotype. VP 9864. Right mandible. 

Locality. Cuba: Matanzas, cave in Sierra de Hato 
Nuevo. March 1917. 
Collector. T. Barbour. 

Condition. Mandible partial — coronoid and angu- 
lar processes broken off. 

Tijpe Series. A series of paratype material is stored 
at the MCZ. 

Comments. G. M. Allen supplemented his descrip- 
tion of Capromys nanus after Thomas Barbour 
provided him with a freshly killed specimen in 
1918 (1918b: 140-145). Type species of the sub- 
genus Paracapromys Kratochvil, Rodriguez, and 
Baiois, 1978. M. nanus was considered a valid spe- 
cies by Woods (1993: 801) and Nowak (1999: 
1703). The type series is stored in the Vertebrate 
Paleontology Department of the MCZ. 

Genus PLAGIODONTA F. Cuvier, 1836 



Plagiodonta araeum Ray, 1 964 
Breviora, Mus. Comp. Zool., 203: 2, 10 
April. 

Holotype. VP 7675. Left upper cheektooth, originally 
described as "almost certainly the fourth (decidu- 
ous) premolar." 

Locality. Dominican Republic: San Rafael Prov- 
ince, Hondo Valle Municipality, unnamed cave 2 
km southeast of Rancho de La Guardia. April 1963. 
Collector R. Allen and C. E. Ray. 
Condition. Condition of tooth as originally de- 
scribed — "damaged slighdy along the posterolabial 
wall." 

Type Series. Holotype only. 

Comments. Considered a valid species by Woods 
(1993: 804) and Nowak (1999: 1708). Complete 
cranial and dentaiy material of araeum has been 
collected in Haiti and deposited in the Florida Mu- 
seum of Natural Histoiy (Woods 1993: 804). This 
species is extinct. This specimen is stored in the 
Vertebrate Paleontology Department of the MCZ. 



Type Specimens of Recent Mammals • Helaen and McFadder 



133 



Order LAGOMORPHA Brandt, 1855 

Family OCHOTONIDAE Thomas, 1897 

Genus OCHOTONA Link, 1795 

Ochotona cuppes Bangs, 1899g 

Proc. New England Zool. Club, 1: 40, 5 

June. 

= Ochotona princeps cuppes Bangs, 1899. 

See A. H. Howell (1924: 27). 

Holotype. B7389. Skin and skull. Adult male. 

Locality. (Canada): British Columbia, Gold Range, 
Monishee Divide, 4,000 ft (1,220 m). 2 August 
1897. 

Collector. A. C. Brooks. Original number 10.30. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; MCZ 7390-7392; all rep- 
resented by sldn and skull, all female. MCZ 7390 
is now at FMNH. 

Comments. O. p. cuppes was retained as a valid 
subspecies by Hall (1981: 289) and Smith and Wes- 
ton (1990: 1). 

Ochotona saxatilis Bangs, 1 899g 

Proc. New England Zool. Club, 1: 41, 5 

June. 

= Ochotona princeps saxatilis Bangs, 

1899. See A. H. Howell (1924: 27). 

Holotype. MCZ 2703. Skin. Adult male. 

Locality. (United States): Colorado, Park County, 
Snowy Range, near Mount Lincoln, Montgomery. 
27 July 1871. 

Collector. J. A. Allen, Rocky Mountain Expedition. 
Original number 945. 
Condition. Skin complete. 

Type Series. Large paratype series; MCZ 209, 243- 
263, skulls; B41 and B42, each represented by sldn 
and skull; MCZ 2673-2703, skins. 
Comments. O. p. saxatilis was retained as a valid 
subspecies by Hall (1981: 291) and Smith and Wes- 
ton (1990: 1). 

Family LEPORIDAE Fischer de Waldheim, 
1817 

Genus LEPL/S Linnaeus, 1758 

Lepus (Macrotolagus) alleni palitans 

Bangs, 1900a 

Proc. New England Zool. Club, 1: 85, 23 

February. 

Holotype. B9096. Skin and skull. Adult female. 

Locality. Mexico: Sinaloa, Aguacaliente (about 40 
miles [64.4 km] southeast of Mazatlan). 7 August 
1897. 



Collector P. O. Simons. Original number 157. 
Condition. Skin and skull complete. Mandible dis- 
articulated; left ramus chipped; two holes in right 
ramus. 

Type Series. 1 paratype; B9097, skin, adult male. 
Comments. Retained as a valid subspecies by Hall 
(1981: 332) and Best and Heniy (1993: 1). 

Lepus americanus struthopus Bangs, 

1898e 

Proc. Biol. Soc. Washington, 12: 81, 24 

March. 

Holotype. B2025. Skin and skull. Adult female. 

Locality. (Canada): Nova Scotia, Digby. 4 August 

1894. 

Collector O. Bangs. Original number 9. 

Condition. Skin and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Comments. Retained as a valid subspecies by Hall 

(1981: 318). 

Lepus arcticus bangsii Rhoads, 1896a 
Amer. Nat., 30: 236, 6 March. 

Holotype. B3752. Skin and skull. Adult female. 

Locality. (Canada): Newfoundland, Codroy. 3 Au- 
gust 1895. 

Collector E. Doane. Original number 1. 
Condition. Sldn and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. Retained as a valid subspecies by Hall 
(1981: 320). 

Lepus bairdii Hayden, 1869 

Amer. Nat., 3: 115, May. 

= Lepus americanus bairdii Hayden, 1869. 

See J. A. Allen (1875: 431). 

Syntype. MCZ 5501. Sldn and skull. Adult. 

Locality. (United States): Wyoming, (Fremont 
County), summit of Wind River Mountains, near 
Fremont Peak. 4 June 1860. 

Collector. F. V. Hayden. Original number 90. For- 
merly USNM 4264. 
Condition. Sldn and skull complete. 
Tijpe Series. 2 other adult syn types; USNM 4262/ 
38001, skin and skull, male; USNM 4263/4273, 
sldn and skidl; see Poole and Schantz (1942: 210). 
Comments. No type is designated in the original 
description, which is based on 6 individuals col- 
lected by Hayden — 3 adults and 3 juveniles, origi- 
nally deposited in the USNM (Poole and Schantz 
1942: 210). 



134 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Lepus bairdi cascadensis Nelson, 1907 

Proc. Biol. Soc. Washington, 20: 87, 11 

December. 

= Lepus americanus cascadensis Nelson, 

1907. See Racey and Cowan (1936: 

HIS). 

Holotype. B1886. Skin and skull. Adult male. 

Locality. Canada: British Columbia, near Hope, 

Roab's Ranch. 12 June 1874. 

Collector. W. C. Colt. Original number 476. 

Condition. Sldn complete. Skull slightly damaged 

(supraoccipital and occipital chipped). 

Type Series. Holotype only. 

Comments. L. a. cascadensis was retained as a valid 

subspecies by Hall (1981: 315). 

Genus Sy/.WMGL/S Gray, 1867 

Lepus (Tapeti) incitatus Bangs, 1901b 
Amer. Nat., 35: 633, 22 August. 
= Sylvilagus brasiliensis incitatus (Bangs, 
1901). See Hershkovitz (1950: 352). 

Holotype. B8441. Skin and skull. Adult female. 

Locality. Panama: (Panama), Bay of Panama, San 

Miguel Island. 30 April 1900. 

Collector W. W. Browi:i, Jr. Original number 127 

Condition. Skin and skull complete. 

Type Series. Holotype only. 

Comments. S. b. incitatus was retained as a valid 

subspecies by Hall (1981: 296). 

Lepus paludicola Miller and Bangs, 1894 

Proc. Biol. Soc. Washington, 9: 105, 9 

June. 

= Sylvilagus palustn's paludicola (Miller 

and Bangs, 1894). See Nelson (1909: 

269). 

Holotype. B1451. Sldn and skull. Adult female. 

Locality. (United States): Florida, (Citrus County), 

Fort Island, near Crystal River. 28 Januaiy 1894. 

Collector F. L. Small. Original number 11.50. 

Condition. Skin and skull complete. 

Type Series. 3 paratypes; B1452, skin and skull, 

adult male; B1453, sldn and skull, adult male; 

B1454, skin and skull, adult female. 

Connnents. S. p. paludicola was retained as a valid 

subspecies by Hall (1981: 299) and Chapman and 

Willner (1981: 1). 

Lepus sylvaticus alacer Bangs, 1 896h 

Proc. Biol. Soc. Washington, 10: 136, 28 

December. 

= Sylvilagus floridanus alacer (Bangs, 

1896). See Lyon (1904: 336). 

Holotype. B5480. Sldn and skull. Adult female. 

Locality. (United States): Indian Territory 



(=Oklahoma), (Adair County), Stilwell. 14 August 
1896. 

Collector. T. Siu-ber. Original number 65. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; B1677, skin and skull, 
B1678, skin and skull, adult male; B5481, skin and 
skull, adult female. 

Comments. S. f. alacer was retained as a valid sub- 
species by Hall (1981: 301) and Chapman et al. 
(1980: 1). 

Lepus sylvaticus transitionalis Bangs, 

1895a 

Proc. Boston Soc. Nat. Hist., 26: 405, 31 

January. 

= Sylvilagus transitionalis (Bangs, 1895). 

See Nelson (1909: 195). 

Holottjpe. B2407. Skin and skull. Adult female. 

Locality. (United States): Connecticut, New Lon- 
don County, Libeity Hill. 6 November 1894. 
Collector O. Bangs. 

Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. There is a series of paratypes in the 
MCZ. 

Comments. S. transitionalis was considered a valid 
species by Hoffman (1993: 827) and Nowak (1999: 
1727). 

Order SCANDENTIA Wagner, 1855 

Family TUPAIIDAE Gray, 1825 

Genus TUPAIA Raffles, 1822 

Tana tana griswoldi CooMge, 1938 

Proc. New England Zool. Club, 17: 45, 6 

May. 

= Tupaia tana paitana (Lyon, 1913). See 

Medway (1965: 76). 

Holotype. MCZ 36416. Sldn, skaill, and baculum. 
Subadult male. 

Locality. (Malaysia): British North Borneo ( = Sa- 
bah). Mount Kinabalu, Kenokok River, Kiau 
( = Kampong Kiau), 3,300 ft (1,007 m). 9 August 
1937. 

Collector J. A. Griswold, Jr., Asiatic Primate Ex- 
pedition. Original number 684. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 



Type Specimens of Recent Mammals • Helpen and McFadden 



135 



Order PRIMATES Linnaeus, 1758 

Family INDRIDAE Burnett, 1828 

Genus PROPITHECUS BenneW, 1832 

Propithecus perrieri Lavauden, 1931 

Compt. Rend. Acad. Sci. Paris, 193: 77, 6 

July. 

= Propithecus diadema perrieri Lavauden, 

1931. See W. C. O. Hill (1953: 568). 

Holotype. MCZ 44857. Skin, skull, and postcranial 
skeleton. Adult male. 

Locality. Malagasy Republic (Madagascar): (Toli- 
aiy), Ifotaka, vicinity of Ambovombe. Probably 
1928. 

Collector. M. Perrier. Original number 1. Grandi- 
dier Collection. 

Conclition. Skin and skull complete. Postcranial 
skeleton partial (right humerus broken, foot bones 
left in sldn). 

Type Series. 3 paratypes; MCZ 44858, skin, skull, 
and postcranial skeleton, adult female; MCZ 
44859, skin and skull, juvenile; MCZ 46001, skin 
and leg bones, adult. 

Comments. P. d. perrieri was retained as a valid 
subspecies by Tattersall (1982: 103). 

Family DAUBENTONIIDAE Gray, 1863 

Genus DAUBENTONIA E. Geoffrey, 1795 

Ctieiromys madagascariensis lanigerG. 

Grandidier, 1930d 

Bull. Acad. Malgache, n. sen, 11: 106 

(for 1928). 

= Daubentonia madagascariensis (Gmelin, 

1788). See G. M. Allen (1939: 134). 

Holotype. MCZ 45947, Skull and postcranial skele- 
ton. Adult male. 

Locality. Madagascar: Forest of the East, 
Collector. Received by G, Grandidier from the 
Academic Malgache, 1927-28. 
Condition. Skull complete. Postcranial skeleton in- 
complete (hands and feet missing, presumably in- 
tact within mounted skin). 
Type Series. Holotype only. 

Comments. The mounted skin of the holotype is in 
the collection of the Academic Malgache (Anta- 
nanarivo Museum). Grandidier named laniger as a 
new subspecies on account of its woolly pelage, but 
it is probably just a molting individual (Schwarz 
1931: 428). 



Family GALAGONIDAE Gray, 1825 

Genus GALAGOIDES A. Smith, 1833 

Galago demidovii orinus Lawrence and 

Wasliburn, 1936 

Occ. Pap. Boston Soc. Nat. Hist., 8: 259, 

8 January. 

= Galagoides orinus (Lawrence and 

Washburn, 1936). See Honess (1996: 58). 

Holotype. MCZ 22453, Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (=Tanzania): Mor- 
ogoro, Uluguru Mountains, Bagilo, 5,000 ft (1,525 
m). 17 September 1926, 
Collector A. Love ridge. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Family GEBIDAE Bonaparte, 1831 

Genus ALOUATTA Lacepede, 1799 

Alouatta palliata luctuosa Lawrence, 

1933b 

Bull. Mus. Comp. Zool., 75: 337, 

November. 

= Alouatta pigra Lawrence, 1933. See 

Smith (1970: 363). 

Holotype. MCZ 24059. Skin and skTill. Adult male. 
Locality. British Honduras ( = Belize): Cayo Dis- 
trict, Mount Cow. 12 April 1928. 
Collector. O. L. Austin, Jr. Original number 723, 
Condition. Skin and skull complete, 
Tijpe Series. HolotyjDe only, 

Alouatta palliata trabeata Lawrence, 

1933b 

Bull. Mus. Comp. Zool., 75: 328, 

November. 

Holotype. MCZ 29545. Skin and skull. Adult male. 
Locality. Panama: Herrera Province, Capina, 
March 1933. 

Collector T Barboui\ Original number 4, 
Condition. Skin and skull complete, 
Tijpe Series. Lawrence mentions examining 19 
specimens of trabeata and lists them by locality 
(1933: 330), 9 of these are in the MCZ; other tlian 
the holotype, they are MCZ 27784, 27785, 28735, 
28736, 29543, 29544, 29546, and 29548, 1 paraty- 
pe, MCZ 29547, has been exchanged to the Museu 
Palista, Sao Paolo, Brazil. 

Comments. Retained as a valid subspecies by Hall 
(1981: 263), 



136 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Genus CEBUS Erxleben, 1777 

Cebus curtus Bangs, 1905b 
Bull. Mus. Comp. ZooL, 46: 91, June. 
= Cebus capucinus curtus Bangs, 1905. 
See Cabrera (1917: 240). 

Holotype. MCZ 10824. Skin and skull. Adult male. 
Locality. Colombia: (Cauca), Gorgona Island. 2 
July 1904. 

Collector. W. W. Brown, Jr. Original number 27. 
Condition. Skin and skull complete. 
Type Series. 1 paratype; MCZ 10825, skin and 
skull, adult female. 

Comments. C. c. curtus was retained as a valid sub- 
species by Cabrera (1958: 169) and Napier (1976: 
36). 

Family HYLOBATIDAE Gray, 1870 

Genus HYLOBATES llliger, 1811 

Hylobates lar carpenteri Groves, 1968 
Proc. Biol. Soc. Washington, 81: 625, 30 
December. 

Holotype. MCZ 41430. Skin and skeleton. Adult 
male. 

Locality. Thailand: Chiengmai District, Mount 
Angka ( = Doi Inthanon), 3,400 ft (1,037 m). 14 
March 1937. 

Collector. H. J. Coolidge, Jr., Asiatic Primate Ex- 
pedition. Original numbers 21 and 185. 
Condition. Skin, skull, and skeleton complete. 
Type Series. Groves examined a series of 144 skins, 
skulls, and skeletons of this subspecies that were 
collected by the Harvard Asiatic Primate Expedi- 
tion in 1937 from Chiengmai District, Thailand, 
"mostly in the Museum of Comparative Zoology, 
Hai'vard, but a few of the osteological specimens 
in the Anthropology Department, University of 
California at Berkeley." He examined also 18 spec- 
imens in the USNM and 1 in the AMNH, all rep- 
resented by skin and skull. 

Comments. Retained as a valid subspecies by Jen- 
kins (1990: 15) and Geissman (1995: 474). 

Family HOMINIDAE Gray, 1825 

Genus GORILLA I. Geoff roy, 1853 

Troglodytes gorilla Savage and Wyman, 

1847 

Boston J. Nat. Hist., 5: 417, December. 

= Gorilla gorilla gorilla (Savage and 

Wyman, 1847). See Rothschild (1923: 

176). 

Lectotype. MCZ 9587. Skull and postcranial skeleton. 
Adult male. 



Locality. Gabon: Gabon Estuaiy, Mpongwe coun- 
tiy. 1847. 

Collector T. S. Savage. Original Jiumber 28 (of J. 
Wyman). 

Condition. Postcranial skeleton partial (includes 
pelvis, sacrum, scapulae, humeri, radii, left ulna, 
femora, left tibia, and 7 vertebrae: 2 cervical, 3 dor- 
sal, 2 lumbar). Head of right humerus and left fe- 
mur bisected. 

Type Series. The original description mentions 
"four skulls, two males and two females, one of 
each in a perfect condition, and all of them adult; 
a male and female peKis, the long bones of the 
upper and lower extremities, and a few vertebrae 
and ribs." However, measurements and illustrations 
of only two of these specimens, an adult male and 
female, each represented by a skull and partial 
postcranial skeleton, are provided. These two, pre- 
sumably the specimens originally described as be- 
ing "in a perfect condition," are the only specimens 
of the original four to be noted in Wymans per- 
sonal notebook of osteology (unpublished, now in 
tlie libraiy of the Boston Museum of Science) and 
die only original specimens of which there is any 
record at all. These two specimens were trans- 
ferred from the Boston Society of Natural History 
to the MCZ in 1915-16. They bear MCZ numbers 
9587 (male) and 9311 (female). The skull of the 
female was sawed in half (hemisected), probably by 
Wyman. The adult male specimen, MCZ 9587, 
possesses an intact skull. MCZ 9587 is the only in- 
tact specimen of the syntype series known to exist, 
and its measurements and an illustration of its skull 
are included in the original publication. Addition- 
ally, it is the only syntype whose measurements are 
included in Coolidge s rex-ision of the genus Gorilla 
(1929: 325), although it is erroneously listed in that 
work as MCZ 9586. For these reasons, MCZ 9587 
is hereby designated as lectotype of Troglodytes go- 
rilla Savage and Wyman, 1847; this should ensure 
taxonomic consistency between past and future 
treatments of this name. 

Comments. Type species of the genus Gorilla I. 
Geoffroy, 1853. The original description is occa- 
sionally attributed in error to Wyman (1847). G. 
gorilla was considered a valid species by Groves 
a993: 276) and Nowak (1999: 618). 

Order LIPOTYPHLA Haeckel, 1866 

Family NESOPHONTIDAE Anthony, 1916 

Genus /VESOPHOA/7ES Anthony, 1916 

Nesophontes micrus G. M. Allen, 1917a 
Bull. Mus. Comp. ZooL, 61: 5, Januaiy. 

Holotype. VP 9600. Right ramus. 

Locality. Cuba: Matanzas Province, cave in Sierra 
de Hato Neuvo. 
Collector C. de la Torre. 



Type Specimens of Recent Mammals • Helgen and McFadden 137 



Condition. Condition as originally described — 
"posterior half of the right ramus, containing a part 
o£ pm^, m„ m.2, and the roots of 111.." 
Type Series. Holotype only. 

Comments. Considered a valid species by Hutterer 
(1993: 70) and Nowak (1999: 201); almost certainly 
extinct. This specimen is stored in the Vertebrate 
Paleontology Department of the MCZ. 

Family SOLENODONTIDAE Gill, 1872 

Genus SOL EA/ODOA/ Brandt, 1833 

Antillogale marcanoi Patterson, 1962 

Breviora, Mus. Comp. ZooL, 165: 3, 22 

August. 

= Solenodon marcano/ (Patterson, 1962). 

See Varona (1974: 8). 

Holotype. VP 7261. Right lower ramus. 

Locality. Dominican Republic: San Rafael Prov- 
ince ( = Elias Pina), Hondo Valle Municipality, un- 
named cave 2 km southeast of Rancho La Guardia. 
Summer 1958. 

Collector. C. E. Ray and A. S. Rand. 
Condition. Condition as originally described — "in- 
complete right ramus of mandible with P3— M, and 
alveoli of other teeth." 

Type Series. 5 paratypes; 7262, left ramus, 7263, 
right humerus, 7264, left humerus, 7265, right 
ulna, 7266, left ramus, juvenile; all partial. 
Comments. A. marcanoi is the type species of the 
genus Antillogale Patterson, 1962. Antillogale was 
first synonymized with Solenodon by Van Valen 
(1967: 255). Considered a valid species by Hutterer 
(1993: 69) and Nowak (1999: 199). S. marcanoi is 
probably extinct. This specimen is stored in the 
Vertebrate Paleontology Department of the MCZ. 

Solenodon poeyanus Barbour, 1944 
Proc. New England Zool. Club, 23: 6, 
March 7. 

= Solenodon cubanus poeyanus Barbour, 
1944. See Aguayo (1950: 131). 

Holotype. MCZ 6597. Mounted skin and skull. 

Locality. Cuba: Oriente ( = Holguin), near Nipi Bay 
( = Nipe Bay). 

Collector Bought by A. Agassiz from H. A. Ward, 
1891. 

Condition. Skin complete. Skull partial (basioccip- 
ital plate and tympanic bullae missing, parietals 
damaged). Mandible disarticulated. 
Type Series. In the original description, Barbour 
includes a photograph of a "living example of So- 
lenodon poeyanus now in Zoological Garden, Ha- 
vana, Cuba, from vicinity of Baracoa." 
Comments. The number of this specimen is 6597, 
not 6957 as stated in the original description. S. c. 



poeyanus was retained as a valid subspecies by Hall 

(1981: 22). 

Family SORICIDAE Fischer von 
Waldheim, 1817 

Genus BLARINA Gray, 1838 

Blarina brevicauda aloga Bangs, 1902a 
Proc. New England Zool. Club, 3: 76, 31 
March. 

Holotype. B9727. Skin and skull. Adult male. 

Locality. (United States): Massachusetts, Dukes 
County, Marthas Vineyard, West Tisbury. 25 June 
1899. 

Collector O. Bangs. Original number 2. 
Condition. Skin and skrdl complete. Mandible dis- 
articulated. 

Type Series. 9 paratypes, B9725, B9726, B9728- 
B9734, all represented by skin and skull, 3 females 
and 6 feinales. 

Comments. Retained as a valid subspecies by Hall 
(1981: 54) and George et al. (1986: 1). 

Blarina brevicauda compacta Bangs, 

1902a 

Proc. New England Zool. Club, 3: 77, 31 

March. 

Holotype. B9705. Skin and skull. Adult male. 

Locality. (United States): Massachusetts, (Nan- 
tucket County), Nantucket (Island). 10 July 1899. 
Collector O. Bangs. Original number 3. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 6 paratypes; B9701-B9704, B9706, 
B9708; all represented by skin and skull, 4 females 
and 2 males. 

Comments. Retained as a valid subspecies by Hall 
(1981: 56) and George et al. (1986: 1). 

Genus CROCIDURAVJagner, 1832 

Crocidura bicolor tephragaster Setzer, 

1956 

Proc. U.S. Nat. Mus., 106: 458, 28 

November. 

= Crocidura fuscomurina (Heuglin, 1865). 

See Hutterer (1983: 223). 

Holotype. MCZ 44773. Skin and skull. Adult male. 
Locality. Anglo-Egyptian Sudan ( = Sudan): (East- 
ern) Equatoria, Torit. 25 April 1950. 
Collector. J. S. Owen. Original number 1158. 
Condition. Sldn and skull complete. 
Type Series. Setzer mentions that he examined 18 
specimens of tephragaster, 8 of which are in the 
MCZ. 



138 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Crocidura hildegardeae phaios Setzer, 

1956 

Proc. U.S. Nat. Mus., 106: 460, 28 

November. 

Holotype. MCZ 45S55. Skin and skull. Adult female. 
Locality. Anglo-Egyptian Sudan ( = Sudan): Equa- 
toria, Imatong Mts, Gilo, 6,500 ft (1,983 m). 12 
June 1950. 

Collector. J. S. Owen. Original number 1266. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; 1 from the MCZ; MCZ 
45856, skin and skull, male. 

Comments. Retained as a valid subspecies, C gra- 
cilipes phaios, by Heim de Balsac and Meester 
(1977: 16); included in Crocidura hildegardeae by 
Hutterer (1993: 87). 

Genus CRYPTOTIS Pomel, 1848 

Cryptotis avia G. M. Allen, 1923a 

Proc. New England Zool. Club, S: 37, 12 

February. 

= Cryptotis thomasi {Mernam, 1897). See 

Woodman (1996: 414). 

Holotype. MCZ 20091. Skin and skull. Adult. 

Locality. Colombia: (probably Cundinamarca), El 
Verjon (see coments). October 1922. 
Collector N. Maria. 

Condition. Sldn complete. Skull partial (most of 
skull from frontals to occiput missing). Left ramus 
of mandible missing. 
Type Series. Holotype only. 

Comments. Considered a valid species by Hutterer 
(1993: 108) and Nowak (1999: 209) but synony- 
mized as noted above. The locality "El Verjon" is 
not shown on any maps available to us but is pos- 
sibly equivalent to "Paramo Cruz Verde" (Paynter 
1997: 463). 

Genus MYOSOREX Gray, 1838 

Crocidura maurisca geata G. M. Allen and 

Loveridge, 1927 

Proc. Boston Soc. Nat. Hist., 38: 417, 23 

December. 

= Myosorex geata (G. M. Allen and 

Loveridge, 1927). See Heim de Balsac 

(1967: 610). 

Holotype. MCZ 22447. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (= Tanzania): Mor- 
ogoro, Uluguru Mountains, Nyingwa, 7,500 ft 
(2,288 m). 19 October 1926. 
Collector A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 



Type Series. 1 paratype; MCZ 22448, skin and 
skull, adult female. 

Comments. M. geata was considered a valid species 
by Hutterer (1993; 99) and Nowak (1999: 217). 

Genus SOREX Linnaeus, 1758 

Neosorex palustris acadicus G. M. Allen, 

1915a 

Proc. Biol. Soc. Washington, 28: 15, 12 

February. 

Name preoccupied by Sorex acadicus 
Gilpin, 1867. 

Sorex palustris gloveralleni Jackson, 1 926 
J. Mammal., 7: 57, 15 February. 
(Replacement name for Neosorex 
palustris acadicus G. M. Allen, 1915) 

Holotype. B2046. Skin and skull. Adult female. 

Locality. (Canada): Nova Scotia, Digby. 26 July 
1894. 

Collector O. Bangs. Original number 3. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B2053, exchanged to the 
USNM in 1922. 

Comments. Retained as a valid subspecies by Hall 
(1981: 41) and Beneski and Stinson (1987: 1). 

Sorex araneus ultimus G. M. Allen, 1914b 

Proc. New England Zool. Club, 5: 51, 9 

April. 

= Sorex tundrensis Merriam, 1900. See 

Hutterer (1993: 121). 

Holotype. MCZ 15000. Skin and skull. Adult male. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth 
of Kolyma River. 6 November 1911. 
Collector. J. Koren. Original number 136. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. There is a series of paratypes, most of 
which are housed in the MCZ. 

Sorex macropygmaeus koreni G. M. Allen, 

1914b 

Proc. New England Zool. Club, 5: 56, 9 

April. 

= Sorex caecutiens koreni G. M. Allen, 

1914. See Yudin (1989: 281). 

Holotype. MCZ 15085. Skin and skiill. Adult female. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth 
of Kolyma River. 19 October 1911. 
Collector J. Koren. Original number .50. 



Type Specimens of Recent Mammals • Helaen and McFadden 



139 



Tijpe Series. 5 paratypes; MCZ 15003-15007, all 
represented by sldn and skull; 4 males, 1 female. 
MCZ 15004, a male, was exchanged to the FMNH 
in 1931. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Sorex macrurus Batchelder, 1896 

Proc. Biol. Soc. Washington, 10: 133, 8 

December. 

Name preoccupied by Sorex macrourus 

Lehmann, 1822. 

Sorex dispar Batchelder, 191 1 
Proc. Biol. Soc. Washington, 24: 97, 15 
May. (Replacement name for Sorex 
macrurus Batchelder, 1896) 

Holotype. MCZ 41744. Skin and skaill. Adult male. 
Locality. (United States): New York, Essex County, 
Keene Heights, Beede's (see comments). 9 Sep- 
tember 1895. 

Collector. C. F. Batchelder. Original number 1384. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; MCZ 41745, skin and 
skull, adult male. 

Comments. S. dispar was considered a valid species 
by Hutterer (1993: 114) and Nowak (1999: 205). 
The type loctility was redescribed by Martin (1966: 
131) as "0.6 mile south and 0.5 mile east of Saint 
Huberts, Essex County, New York, lat. 44°09', long. 
73°46'." 

Sorex personatus miscix Bangs, 1899d 

Proc. New England Zool. Club, 1: 15, 28 

Februaiy. 

= Sorex cinereus miscix Bangs, 1899. See 

Jackson (1925: 56). 

Holotype. B8651. Skin and skull. Adult male. 

Locality. (Canada): Labrador, Black Bay. 10 Octo- 
ber 1898. 

Collector E. Doane. Original number 1. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. Bangs based his description on 39 
specimens; corresponding to B7931-B7950 and 
B8651-B8669; all represented by skin and skull. 
Comments. S. c. miscix was retained as a valid sub- 
species by Hall (1981: 29). 

Sorex sanguinidens G. M. Allen, 1914b 

Proc. New England Zool. Club, 5: 54, 9 

April. 

= Sorex daptiaenodon sanguinidens G. M. 

Allen, 1914. See Yudin (1989: 198). 

Holotype. MCZ 15012. Skin and skull. Adult female. 
Locality. (Russian Federation): northeastern Sibe- 



ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth 

Kolyma River. 11 December 1911. 

Collector J. Koren. Original number 221. 

Condition. Tail separate from sldn. Skull complete. 

Mandible disarticulated. 

Type Series. There is a large series of paratypes, 

most of which are still in the MCZ. 

Sorex virG. M. Allen, 1914b 

Proc. New England Zool. Club, 5: 52, 9 

April. 

= Sorex roboratus Hollister, 1913. See 

Hoffman (1985: 17). 

Holotype. MCZ 15068. Skin and skull. Adult female. 
Locality. (Russian Federation): northeastern Sibe- 
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth 
of Kolyma River. 19 December 1911. 
Collector J. Koren. Original number 230. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a large series of paratypes, 
most of which are still in the MCZ. 

Genus SL/A/CL/S Ehrenberg, 1832 

Suncus aterMedway, 1965 

Mammals of Borneo, J. Malay. Branch R. 

Asiat. Soc, 36: 38, December. 

Holotype. MCZ 36574. Skin, skull, and postcranial 
skeleton. Adult female. 

Locality. Malaysia: northern Borneo, Sabah, 
Mount Kinabalu, Lumu Lumu, 5,500 ft (1,678 m). 
7 July 1937. 

Collector J. A. Griswold, Jr. Original number 462. 
Condition. Skin complete. Skull partial (tympanic 
bullae missing), and mandible disarticulated. Post- 
cranial skeleton complete. 
Type Series. Holotype only. 

Comments. Considered a valid species by Hutterer 
(1993: 101) and Nowak (1999: 223). 

Suncus varilla minor G. M. Allen and 

Loveridge, 1933 

Bull. Mus. Comp. Zool, 75: 57, 

Februaiy. 

Holotype. MCZ 26754. Skin and skaill. Adult female. 
Locality. Tanganyika Territory (=Tanzania): Urun- 
gu, Kitungulu, 4,500 ft (1,373 m). 14 May 1930. 
Collector. A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotyj^e only. 

Comments. Retained as a valid subspecies by Heim 
de Balsac and Meester (1977: 6). 



140 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Family TALPIDAE Fischer de Waldheim, 
1817 

Genus NEUROTRICHUS Gm\her, 1880 

Neurotrichus gibbsi hyacinthinus Bangs, 

1897d 

Amer. Nat., 31: 240, 1 March. 

Holotype. B1240. Sldn and skull. Adult female. 

Locality. U.S.A. (United States): California, Marin 
County, Nicasio. 10 March 1894. 
Collector. C. A. Allen. Original number 694. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B1241, skin and skull, 
adult male. 

Comments. Retained as a valid subspecies by Hall 
(1981: 67) and Carraway and Verts (1991: 1). 

Genus SCALOPUS Desmarest, 1 804 

Scalops anastasae Bangs, 1898b 

Proc. Boston Soc. Nat. Hist., 28: 212, 15 

March. 

= Scalopus aquaticus anastasae (Bangs, 

1898). See Jackson (1915: 39). 

Holotype. B7192. Skin and skull. Adult male. 

Locality. U.S.A. (United States): Florida, St. Johns 
County, Anastasia Island, Point Romo. 16 Febioiary 
1897. 

Collector O. Bangs. Original number 10. 
Condition. Skin and skaiU complete. Mandible dis- 
articulated. 

Tijpe Series. 4 paratypes; B719.3-B7196; all repre- 
sented by sldn and skull, 2 females and 2 males. 
Comments. S. a. anastasae was retained as a valid 
subspecies by Hall (1981: 72). 

Scalops texanus aereus Bangs, 1896li 

Proc. Biol. Soc. Washington, 10: 138, 28 

December. 

= Scalopus aquaticus aereus (Bangs, 

1896). See Miller (1912: 8). 

Holotype. B547.5. Skin and skull. Adult female. 

Locality. (United States): Indian Territoiy 
( = Oklalioma) (Adair County), Stilwell. 13 August 
1896. 

Collector. T. Surber. Original number 64. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. S. a. aereus was retained as a valid sub- 
species by Hall (1981: 72). 



Genus SCAPANUS Pome\, 1848 

Scapanus californicus minusculus Bangs, 

1899J 

Proc. New England Zool. Club, 1: 70, 31 

Jnly. 

= Scapanus latimanus minusculus Bangs, 

1899. See Grinnell and Swarth (1912: 

133). 

Holotype. B9189. Skin and skull. Adult female. 
Locality. (United States): California, El Dorado 
County, Fyffe. 10 June 1897. 

Collector W. W. Price and E. M. Nutting. Original 
number 15. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. S. I. minusculus was retained as a valid 
subspecies by Hall (1981: 70). 

Order CHIROPTERA Blumenbach, 1779 

Family PTEROPODIDAE Gray, 1821 

Genus AETHALOPS Thomas, 1923 

Aethalops aequalis G. M. Allen, 1938b 
J. Mammal., 19: 497, 14 November. 
= Aethalops alecto aequalis G. M. Allen, 
1938. See Hill (1961: 639). 

Holotype. MCZ 36582. Skin, skull, and postcranial 
skeleton. Adult female. 

Locality. (Malaysia): British North Borneo ( = Sa- 
bali). Mount Kinabalu, Luma Luma, 5,500 ft (1,678 
m). 12 July 1937. 

Collector. J. A. Griswold, Jr., Asiatic Primate E.x;- 
pedition. Original number 510. 
Condition. Sldn, skull, and postcranial skeleton 
complete. 

Type Senes. 3 paratypes; MCZ 36583, sldn, skull 
and skeleton, female; MCZ 36584, skin and skull, 
female; MCZ 36586, skin and skull, female. 
Comments. A. a. aequalis was retained as a valid 
subspecies by Koopman (1994: 35). 

Genus EONYCTERIS Dobson, 1873 

Eonycteris spelaea glandifera Lawrence, 

1939 

7/7 Barbour, Lawrence, and Peters, Bull. 

Mus. Comp. Zool., 86: 38, November. 

Holotype. MCZ 35159. Sldn and skull. Adult male. 
Locality. Philippines: Luzon, (Nueva Ecija), Rizal, 
Montalban caves near Manila. 27 Febiniary 1937. 
Collector. B. Lawrence. Original number 253. 
Condition. Skin and skull complete. 
Type Series. In the original description, Lawrence 



Type Specimens of Recent Mammals • Helaen and McFadden 



141 



implies the existence of several specimens in ad- 
dition to the holotype; she collected 18 specimens 
of glandifera during her 1936—37 expedition to the 
Philippines, all of which are in the MCZ. 
Comments. Retained as a valid subspecies by Ma- 
haradatunkamsi and Kitchener (1997: 59). 

Genus HAPLONYCTERIS Lawrence, 
1939 

Haplonycteris fischeri Lawrence, 1939 
In Barbour, Lawrence, and Peters, BulL 
Mus. Comp. ZooL, 86: 33, November. 

Holotype. MCZ 35258. Skin and skull. Adult male. 
Locality. Philippines: Mindoro (Oriental), Mouirt 
Halcon, Bignay. 26 April 1937. 
Collector F. S. Rivera. Original number BL 502. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. Type species of the genus Haplonyc- 
teris Lawrence, 1939. Considered a valid species 
by Koopman (1993: 142) and Nowak (1999: 292). 

Genus PTEROPUS Erxleben, Mil 

Pteropus anetianus aorensis Lawrence, 

1945 

Proc. New England ZooL Club, 23: 66, 

26 March. 

Holotype. MCZ 42183. Skin and skull. Adult male. 
Locality. (Vanuatu): New Hebrides, off southwest 
corner of Espiritu Santo Island, Aore Island. 8 
April 1944. 

Collector O. L. Austin, Jr. Original number 5. 
Condition. Skin complete. Skull partial (basioccip- 
ital missing), right ramus of mandible broken. 
Type Series. 1 paratype; MCZ 42182, skin and 
skull, adult male. 

Comments. Retained as a valid subspecies by 
Koopman (1994: 25) and Flanneiy (1995b). 

Pteropus arielG. M. Allen, 1908 
Bull. Mus. Comp. ZooL, 52: 28, July. 
= Pteropus giganteus ariel G. M. Allen, 
1908. See Hill (1958: 5). 

Holotype. MCZ 10565. Skin and skull. Adult male. 
Locality. Maldive Islands: Male Atoll. 24 Decem- 
ber 1901. 

Collector. H. B. Bigelow, A. Agassiz E.xpedition. 
Condition. Right wing of sldn damaged. Skull in- 
tact, with 2 small holes in braincase. 
Type Series. 1 paratype; MCZ 10566, skin and 
skull, juvenile female. 

Comments. P. g. ariel was retained as a valid sub- 
species by Koopman (1994: 26). 



Pteropus austini Lawrence, 1945 

Proc. New England Zool. Club, 23: 59, 

26 March. 

= Pteropus woodfordi Thomas, 1888. See 

Sanborn and Beecher (1947: 389). 

Holotype. MCZ 42166. Skin and skull. Subadult fe- 
male. 

Locality. Solomon Islands: Florida Island (Nggela 
Group). 20 Februaiy 1944. 
Collector O. L. Austin, Jr. Original number 2. 
Condition. Skin and skull complete. 
Type Series. 1 paratype; MCZ 42167, skin and 
skull, subadult male. 

Comments. Lawrence referred to the holotype as 
an adult in the original description, but Sanborn 
and Beecher (1947: 389) recognized it as a sub- 
adult. 

Pteropus rayneri monoensis Lawrence, 

1945 

Proc. New England Zool. Club., 23: 63, 

26 March. 

Holotype. MCZ 42191. Skin and skull. Aduh male. 
Locality. Solomon Islands: Treasuiy (Mono) Island. 
11 October 1944. 

Collector O. L. Austin, Jr. Original number 27. 
Condition. Skin and skull complete. 
Type Series. 2 paratypes; MCZ 42192, skin and 
skull, adult male; MCZ 42193, skin and skull, adult 
male. 

Comments. Retained as a valid subspecies by 
Koopman (1994: 24) and Flannery (1995b: 285). 

Genus ROUSETTUS Gray, 1821 

Rousettus madagascariensis G. 

Grandidier, 1930b 

Bull. Acad. Malgache, n. sen, 11: 91 (for 

1928). 

Holotype. MCZ 45432. Alcoholic and skull. Adult 
male. 

Locality. Malagasy Republic (Madagascar): (Anta- 
nanarivo), between Tananarive ( = Antananarivo) 
and Andevoranto, Grand forest de Est, near Be- 
forona. 

Collector Received by G. Grandidier from the 
Academic Malgache, 1917. 
Conditio72. Alcoholic, skull complete. 
Type Series. Holotype only. 

Comments. Considered a valid species by Koop- 
man (1993: 153) and Nowak (1999: 261). In his 
review of known material of jR. madagascariensis, 
Bergmans (1977: 67) commented in error that the 
holotype was in the Academic Malgache, Antana- 
narivo, Madagascar. 



142 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Family EMBALLONURIDAE Gervais, 1855 

Genus PEROPTERYX Peters, 1867 

Peropteryx canina phaea G. M. Allen, 

1911a 

Bull. Mus. Comp. ZooL, 54: 222, July. 

= Peropteryx macrotis phaea G. M. Allen, 

1911. See Sanborn (1937: 342). 

Holotijpe. MCZ 8101. Skin and skull. Adult female. 
Locality. Grenada: Point Saline(s). 29 August 1910. 
Collector. G. M. Allen. Original number 15. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. There is a series of paratypes in the 
MCZ. 

Comments. P. m. phaea was retained as a valid sub- 
species by Hall (1981: 82) and Koopman (1994: 
47). 

Genus RHYNCHONYCTERIS Peters, 
1867 

Rhynchiscus naso priscus G. M. Allen, 

1914c 

Proc. Biol. Soc. Washins^ton, 27: 109, 10 

July. 

= Rhynchonycteris naso (Wied-Neuwied, 

1820). See Sanborn (1937: 326). 

Holotijpe. MCZ 13208. Skin and skull. Adult. 

Locality. Me.\ico: Quintana Roo, Xcopen. 18 Feb- 
ruary 1912. 

Collector. J. L. Peters. Original number 13. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; MCZ 13209, skin and 
skuU, adult male; MCZ 14637, alcoholic, adult 
male; MCZ 14638, alcoholic, adult female. 

Family NYCTERIDAE Van der Hoeven, 
1855 

Genus NYCTERIS G. Cuvier and E. 
Geoffroy, 1795 

Nycteris madagascariensis G. Grandidier, 

1937 

Bull. Mus. Nat. Hist. Paris, 9: 353, 25 

November. 

Holotype. MCZ 45433. Body in alcohol skull extract- 
ed. 

Locality. Madagascar: (Antsiranana), Diego-Suarez 
( = Antsiranana), Valley of the Rodo, north of Pir- 
kana near the Ankarana, 12°5'-13°0'S, 49°5'E. June 
1910. 
Collector. Grandidier collection. 



Condition. Alcoholic, skull complete. 

Type Series. 1 paratype, MCZ 45434, in alcohol, 

skull extracted, female. 

Comments. N. madagascariensis was included in N. 

macrotis by Koopman (1993: 162) but retained as 

a valid species by Peterson et al. (1995; 6.3). 

Nycteris nana tristis G. W\. Allen and 
Lawrence, 1936 

Bull. Mus. Conip. Zool., 79: 47, January. 
^Nycteris nana (Andersen, 1912). See 
Hayman and Hill (1971: 19). 

Holotype. MCZ 31156. Skin and skull. Adult female. 
Locality. Kenya: (W. Nyanza), Kakamega District, 
Kaimosi. 13 Februaiy 1934. 
Collector A. Loveridge. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 
Type Series. Holotype only. 

Family RHINOLOPHIDAE Gray, 1825 

Genus HIPPOSIDEROSGmy, 1831 

Hipposideros curtus G. M. Allen, 1921 
Rev. Zool. Africaine, 9: 194, December. 

Holotype. MCZ 19305. Body in alcohol. Female. 
Locality. Cameroons ( = Cameroon): (Littoral), Sak- 
bayeme. 1920. 
Collector G. Schwab. 

Condition. Alcoholic. Skull extracted but not 
cleaned (still in alcohol). 
Type Series. Holotype only. 

Comments. H. curtus was considered a valid spe- 
cies by Koopman (1993: 172) and Nowak (1999: 
333). ' 

Hipposideros erigens Lawrence, 1939 
In Barbour, Lawrence, and Peters, Bull. 
Mus. Comp. Zool., 86: 56, November. 
= l-lipposideros bicolor erigens Lawrence, 
1939. See Hill (1963: 28). 

Holotype. MCZ 35197. Skin and skull. Adult male. 
Locality. Philippines: Mindoro, (Oriental) Tabucala 
cave near Calapan, northern base of Mount Hal- 
con. 7 March 1937. 

Collector B. Lawrence. Original number 307. 
Condition. Skin complete. Skull partial (occiput 
missing), and mandible disarticulated. 
Type Series. 3 paratypes; MCZ 35195, skin and 
skull, adult female; MCZ 35196, sldn and skull, 
adult male; MCZ 35198, skin and skull, adult fe- 
male. 

Comments. H. b. erigens was retained as a valid 
subspecies by Koopman (1994; 61). 



Type Specimens of Recent Mammals • Helaen and McFadden 



143 



Hipposideros turpis Bangs, 1901a 
Amer. Nat., 35: 561, 31 July. 
= Hipposideros turpis turpis Bangs, 1901. 
See Hill (1963: 94). 

Holotijpe. MCZ 10003. Skin and skull. Adult female. 
Locoliti/. (Japan): Ryaikyu Islands, southern group 
of Liu kin Islands, Ishigaki Island. 10 May 1899. 
Collector. I. Zensaku. 

Condition. Sldn complete. Skull partial (occipital 
region missing). 

Type Series. 2 paratypes; MCZ 10002, skin and 
skull, adult female; MCZ 10004, skin and skvill, 
adult male. 

Comments. H. tuiyis was considered a valid species 
by Koopman (1993: 175) and Nowak (1999: 334). 

Genus RHINOLOPHUS Lacepe6e, 1799 

Riiinoloplius megaphyllus igniter G. M. 

Allen, 1933 

J. Mammal., 14: 149, 15 May. 

= Rtiinolophus megapfiyilus megapiiyllus 

Gray, 1834. See Koopman (1984: 9). 

Holotijpe. MCZ 29078. Skin and skull. Adult male. 
Locality. Australia: Queensland, Cape York, Coen. 
12 June 1932. 

Collector P. J. Darlington, Jr., Hai-vard Australian 
Expedition. Original number 185. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratyj^jc; MCZ 29079, skin and 
skull, adult male. 

Rtiinolophus ptiilippinensis alleni 
Lawrence, 1939 

In Barbour, Lawrence, and Peters, Bull. 
Mus. Comp. Zool., 86: 46, November. 

Holotype. MCZ 35097. Sldn and skull. Adult female. 
Locality. Philippines: Mindoro, (Oriental) Tabucala 
cave near Calapan, northern base of Mount Hal- 
con. 7 March 1937. 

Collector. B. Lawrence. Original number 302. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 2 pai'atypes; MCZ 35098, skin and 
sk-ull, adult female; MCZ 35099, skin and skull, 
adult female. 

Comments. Retained as a valid subspecies by 
Koopman (1994: 57). 

Genus TRIAENOPS Dobson, 1871 

Triaenops aurita G. Grandidier, 1912 
Bull. Mus. Hist. Nat. Paris, 18: 8, 25 
Januaiy. 

Holotype. MCZ 45080. Mummy. 

Locality. Malagasy Republic ( = Madagascar): (An- 
tsiranana), Diego-Suarez (=Antsiranana). 1910. 



Collector. Dr. Mazieres. 

Condition. Mummy (dried carcass), complete. 
Type Series. Holotype only. 

Comments. Traditionally included in Triaenops fur- 
culus, as in G. M. Allen (1939: 82) and Koopman 
(1993: 175), but retained as distinct by Peterson et 
al. (1995: 81) pending further material from the 
area of the type locality. Known only from the ho- 
lotype. 

Family MORMOOPIDAE de Saussure, 
1860 

Genus PTERONOTUS Gray, 1838 

Ctiilonycteris parnellii pusillus G. M. Allen, 

1917c 

Proc. Biol. Soc. Washington, 30: 168, 23 

October. 

= Pteronotus parnellii pusillus (G. M. Allen, 

1917). See Smith (1972: 67). 

Holotype. MCZ 16468. Skin and skull. Female. 
Locality. Dominican Republic: Santo Domingo, 
Arroyo Salado. 7 March 1916. 
Collector J. L. Peters. Original number 227. 
Condition. Skin complete. Skull partial (left wall of 
braincase broken, left tympanic bulla missing). 
Type Series. 2 paratypes; MCZ 16599, female; 
MCZ 16600, female; both in alcohol. 
Comments. P. p. pusillus was retained as a valid 
subspecies by Hall (1981: 92) and Koopman (1994: 
71). 

Ctiilonycteris torrelG. M. Allen, 1916a 

Proc. New England Zool. Club, 6: 4, 8 

Februai-v. 

= Pteronotus quadridens quadridens 

(Gundlach, 1840). See Silva-Taboada 

(1976: 7). 

Holotype. MCZ 11672. Body in alcohol, skull extract- 
ed. Adult female. 

Locality. Cuba: (Guantanama), Baracoa, La Cueva 
de la Majana. 15 June 1915. 

Collector V. J. R. Verrier. Presented to the MCZ 
by Carlos de la Torre. 
Condition. Alcoholic, skull complete. 
Type Series. 2 paratypes; MCZ 11670, male; MCZ 
11671, female; both in alcohol. 

Family PHYLLOSTOMIDAE Gray, 1825 

Genus AMETRIDA Gray, 1847 

Ametrida minor H. Allen, 1894 

Proc. Boston Soc. Nat. Hist., 26: 240, 16 

May. 

= Ametrida centurio Gray, 1847. See 

Peterson (1965: 5). 

Holotype. MCZ 11274. Body in alcohol, skull extract- 
ed. Adult male. 



144 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Locality. Suriname: Paramaribo. Collected some- 
time between 1832 and 1839. 
Collector. F. W. Cragin. 
Condition. Alcoholic, skull complete. 
Type Series. Holotype only. 

Comments. For comments on the type locality and 
date of acquisition, see G. M. Allen (1902a: 88). A. 
minor actually represents the male specimens of A. 
centurio. 

Genus ARTIBEUS Leach, 1821 

Artibeus femurvillosum Bangs, 1899k 

Proc. New England Zool. Club, 1: 73, 24 

November. 

= Artibeus lituratus palmarum J. A. Allen 

and Chapman, 1897. See Hershkovitz 

(1949: 445). 

Holotype. B8314. Skin and skull. Adult male. 

Locality. Colombia: (La Guajira), La Concepcion, 

3,000 ft (915 m). 21 March 1899. 

Collector W. W. Brown, Jr. Original number 31. 

Condition. Skin and skull complete. 

Ti/pe Series. Holotype only. 

Genus EROPHYLLA Miller, 1906 

Erophylla sezekomi syops G. M. Allen, 

1917c 

Proc. Biol. Soc. Washington, 30: 167, 23 

October. 

Holotype. MCTj 13713. Body in alcohol, skull extract- 
ed. Adult male. 

Locality. Jamaica: (St. James), Montego Bay. 14 
March 1912. 

Collector J. A. Cushman. 
Condition. Alcoholic, skull complete. 
Type Series. 7 paratypes; MCZ 13709-13712, 
13714-13716, all alcoholic. MCZ 13709 and 13712 
were sent in e.xchange to the USNM. 
Comments. Retained as a valid subspecies by Hall 
(1981: 171) and Koopman (1994: 79). 

Genus GLOSSOPHAGA E. Geoffrey, 
1818 

Glossophaga longirostris Miller, 1898 
Proc. Acad. Nat. Sci. Philadelj^hia, 1898, 
p. 330, 2 August. 

= Glossophaga longirostris longirostris 
Miller, 1898. See Miller (1913: 422). 

Holotype. B8046. Skin and skull. Adult female. 

Locality. Colombia: (Magdalena), Santa Marta 
Mountains, near Santa Marta. 10 February 1898. 
Collector W. W. Brown, Jr. Original number 60. 



Condition. Skin complete. Skull partial (left zygo- 
matic arch missing). 
Type Series. Holotype only. 

Comments. G. longirostris was considered a valid 
species by Koopman (1993: 184) and Nowak (1999: 
368). 

Genus LONCHOPHYLLA Thomas, 1903 

Lonchophylla hesperia G. M. Allen, 1908 
Bull. Mus. Comp. Zool, 52: 35, July. 

Holotype. MCZ 7011. Body in alcohol, skull extract- 
ed. Adult male. 

Locality. Peru: (Contralmirante Villar), Tumbes, 
Zorritos. 

Collector. F. H. Bradley. 
Condition. Alcoholic, skull complete. 
Type Series. 2 paratypes; in the YPM; 1034 and 
1035; both in alcohol. 

Comments. Considered a valid species by Koop- 
man (1993: 181) and Nowak (1999: 372). This rare- 
ly collected bat is known by only two museum 
specimens in addition to the type series: USNM 
283177 and LSUMZ 14121 (Gardner 1976: 5). 

Genus PLATYRRHINUS 6e Saussure, 
1860 

Vampyrops umbratus Lyon, 1902 

Proc. Biol. Soc. Washington, 15: 151, 20 

June. 

= Platyrrhinus umbratus (Lyon, 1902). See 

Koopman (1993: 191). 

Holotype. B8180. Sldn and skull. Adult male. 

Locality. Colombia: (LaGuajira), San Miguel. 8 
June 1898. 

Collector W. W. Brown, Jr. Original number 234. 
Condition. Sldn and skull complete. 
Type Series. 2 paratypes; B8300, skin, male; B8301, 
sldn, male. 

Comments. P. umbratus was considered a valid spe- 
cies by Koopman (1993: 191) and Nowak (1999: 
389). Platyrrhinus has priority over the genus name 
Vampyrops (Gardner and Ferrell 1990: 501-503). 

Vampyrops zarhinus H. Allen, 1891 
Proc. Acad. Nat. Sci. Philadelphia, 1891, 
p. 400, 22 September. 
= Platyrrhinus helleri {Peters, 1866). See 
Hall and Kelson (1959: 131). 

Holotype. MCZ 3211. Body in alcohol, skull extract- 
ed. Adult female, pregnant. 

Locality. Panama: Canal Zone, Obispo. 1872. See 
comments. 

Collector Hassler Expedition. 
Condition. Alcoholic, skull complete. 
Type Series. Holotype only. 



Type Specimens of Recent Mammals • Helgen and McFadden 145 



Comments. In the original description, H. Allen re- 
port:ed that this specimen had been collected in 
Brazil by the Thayer expedition. G. M. Allen 
emended this apparently erroneous locality to 
Obispo, Panama, in accordance ^A-ith the accession 
catalogue of the MCZ (1931: 236-237). In support 
of Aliens decision, Rouk and Carter (1972: 4) stat- 
ed, after examining the holotype of zarhinus, that 
it is "quite like specimens of [Platyrrhiniis] helleri 
from Mexico and Central America, and unlikely to 
have come from Brazil." For the use of the genus 
Platyrrhinus over Vampyrops, see Gardner and 
Ferrell (1990: 501-503). 



Genus VAMPYRODES Thomas, 1 900 

Vampyrodes major G. M. Allen, 1908 
Bull. Mus. Comp. ZooL, 52: 38, July. 
= Vampyrodes caraccioli major G. M. 
Allen, 1908. See Handley (1966: 766). 



Holotype. MCZ 6756. Body in alcohol. Adult female. 
Locality. Panama: San Pablo (now covered by Ga- 
tun Lake). Date unrecorded. 
Collector. A. Lesley. 
Condition. Alcoholic. 
Type Series. Holotype only. 

Comments. V. c. major was retained as a valid sub- 
species by Koopman (1994: 88). 



Family MOLOSSIDAE Gervais, 1855 

Genus MOPS Lesson, 1842 

Chaerephon leucostigma G. M. Allen, 

1918a 

Bull. Mus. Comp. ZooL, 61: 513, 

Februaiy. 

= Mops condylurus leucostigma (G. M. 

Allen, 1918). See Koopman (1994: 141). 



Holotype. MCZ 16344. Skin and skull. Aduh female. 
Locality. Malagasy Republic (Madagascar): (Anta- 
nanarivo), Tananarive (= Antananarivo). December 
1915. 

Collector. F. R. Wulsin. 

Condition. Skin partial (bare spot on ventrum). 
Skull damaged (right and left zygomatic arch miss- 
ing; supraoccipital chipped). 

Type Series. 1 paratype; MCZ 16345, skin and 
skull, male. 

Comments. Peterson et al. (1995: 168) used the 
name Tadarida leucostigma. 



Mops angolensis orientis G. M. Allen and 

Loveridge, 1942 

Bull. Mus. Comp. ZooL, 89: 166, 

Februaiy. 

= Mops condylurus orientis G. M. Allen 

and Loveridge, 1942. See Koopman 

(1994: 141). 

Holotype. MCZ 38829. Skin and skull. Adult male. 
Locality. Tanganyika Territoiy (= Tanzania): Mtwa- 
ra, Ruviuna River, Kitaya, 300 ft (92 m). 3 April 
1939. 

Collector A. Loveridge. 
Condition. Skin and skull complete. 
Tijpe Series. 9 paratypes; MCZ 38826-38828, 
38830-38835, all represented by skin and skull, 4 
females and 5 males. 

Genus OTOMOPS Thomas, 1913 

Otomops papuensis Lawrence, 1948 
J. Mammal., 29: 413, 31 December. 

Holotype. MCZ 45769. Body in alcohol, skull extract- 
ed. Adult female. 

Locality. Papua New Guinea: Vailala River. 
Collector. Bought from Ward's Natural Science Es- 
tabhshment, April 1948. 

Condition. Alcoholic, skull partial (right zygomatic 
arch missing). 
Type Series. Holotype only. 

Comments. Considered a valid species by Koop- 
man (1993: 239) and Nowak (1999: 482). Accord- 
ing to Flanneiy (1995a: 481), O. papuensis has 
been collected on only two occasions and, other 
than the holotype, is known by only 10 specimens; 
2 in the BMNH, the remainder in the biological 
collections of the University of Papua New Guinea. 

Family VESPERTILIONIDAE Gray, 1821 

Genus EPTES/CL/S Rafinesque, 1820 

Eptesicus darlingtoni G. M. Allen, 1933 
J. Mammal., 14: 150, 15 May. 

Holotype. MCZ 29113. Skin and skull. Adult female. 
Locality. Australia: Queensland, Queensland Na- 
tional Park, MacPherson Ranges, 3,000 ft (915 m). 
10 March 1932. 

Collector P. J. Darlington, Jr., Harvard Australiaji 
Expedition. Original number 30. 
Condition. Skin and skull complete. 
Type Series. 1 paratype; MCZ 29120 (now Queens- 
land Museum J 5476), skin and skull, adult female. 
Comments. McKean et al. (1978: 533) and Koop- 
man (1993: 203) included darlingtoni in Eptesicus 
pumilus. However, Koopman also used the name 
Pipistrellus darlingtoni (1994: 116). Hoye (1995: 



146 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



537) considered darlingtoni to be a valid species of 
VespadeJus, to which he gave full generic rank. 

Genus HARPIOCEPHALUS Gray, 1842 

Harpiocephalus rufulus G. M. Allen, 1913 
Proc. Biol. Soc. Washington, 26: 214, 20 
December. 

= Harpiocephalus harpia rufulus G. M. 
Allen, 1913. See Ellerman and Morrison- 
Scott (1951: 187). 

Holotijpe. MCZ 14206. Skin and skull. Adult male. 
Localitij. Vietnam: Tonkin, Lao-Kai ( = Lao Cai). 3 
Januaiy 1912. 

Collector. Kobayashi Collection. Original number 
14. 

Condition. Skin complete. Skull partial (parietals 
broken). 

Type Serie.s. Holotype only. 

Comments. H. h. nifuliis was retained as a valid 
subspecies by Koopman (1994: 133). 

Genus IDIONYCTERIS Anthony, 1923 

Corynorhinus phyllotis G. M. Allen, 1916b 
Bull. Mns. Comp. Zool., 60: 352, April. 
= Idionycteris phyllotis phyllotis (G. M. 
Allen, 1916). See Tumlison (1993: 418). 

Holotype. MCZ 5943. Skin and skull. Adult. 

Locality. Mexico: San Luis Potosi. 24 March 1878. 
Collector. E. Paliner. 
Condition. Skin and skull complete. 
Tijpe Series. Holotype only. 

Comments. I. phyllotis was considered a valid spe- 
cies by Koopman (1993: 205) and Nowak (1999: 
457). 

Genus LASIURUS Gray, 1838 

Atalapha brachyotisJ. A. Allen, 1892 

Bull. Amer. Mus. Nat. Hist., 4: 47, 25 

March. 

= Lasiurus borealis brachyotis (J. A. Allen, 

1892). See Niethammer (1964: 595). 

Holotype. MCZ 11143. Body in alcohol. Male. 

Locality. (Ecuador), Galapagos Islands: Chatham 
Island. 23 June 1891. 
Collector G. Baur. 

Condition. Alcoholic. The specimen was received 
without a skull. 
Type Series. Holotype only. 

Comments. Lasiunis brachyotis has often been ac- 
corded specific status, as in Nowak (1999: 451). In- 
cluded in L. borealis as a valid subspecies by Koop- 
man (1994: 129). 



Genus /W/OT/S Kaup, 1829 

Myotis abbotti nugax G. M. Allen and 

Coolidge, 1940 

Bull. Mus. Comp. Zool., 87: 137, 31 

December. 

= Myotis muricola nugax G. M. Allen and 

Coolidge, 1940. See Koopman (1994: 

104). 

Holotype. MCZ 36076. Skin and skull. Adult male. 
Locality. Malaysia: north Borneo, Sabah, Mount 
Kinabalu, Bundutuan, 3,500 ft (1,068 m). 25 July 
1937. 

Collector J. A. Griswold, Jr., Asiatic Primate Ex- 
pedition. Original number 626. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 16 paratypes, MCZ 36072-36075, 
36077-36080, 36082-83, 36085-89, 36091; all rep- 
resented by skin and skull, 12 females and 4 males. 

Myotis albicinctus G. M. Allen, 1919b 
J. Mammal., 1: 2, 28 November. 
= Myotis lucifugus carissima Thomas, 
1904. See Miller and G. M. Allen (1928: 

50). 

Holotype. MCZ 11747. Sldn and skull. Adult male. 
Locality. (United States): California, (Tulare Coun- 
ty), Mount Whitney, 11,000 ft (3,355 m). 14 July 
1915. 

Collector G. M. Allen. Original number 1. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratyj^^e, probably at USNM. 
Comments. The skull of the holotype, which had 
been mislaid at the time that albicinctus was de- 
scribed, has subsequently been found and reunited 
with its skin. 

Myotis sodalis Miller and G. M. Allen, 

1928 

Bull. U.S. Nat. Mus., 144: 130, 25 May. 

Holotype. MCZ 10988. Skin and skull. Adult female. 
Locality. (United States): Indiana, (Crawford 
County), Wyandotte cave. 7 March 1904. 
Collector J. O. Sibert. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Comments. Considered a valid species by Koop- 
man (1993: 215) and Nowak (1999: 419). 
Type Series. Miller and Allen based their descrip- 
tion on an examination of 443 specimens, described 
by locality in the original description (1928: 133). 
30 paratypes are in the MCZ, the others are in tlie 
FMNH, USNM, AMNH. and BMNH. 



Type Specimens of Recent Mammals • Hehen and McFadden 147 



Genus A/VCT/CE/L/S Rafinesque, 1819 

Nycticeius africanus G. M. Allen, 1911b 

Bull. Mus. Comp. Zool., 54: 328, 

December. 

= Nycticeius sctilieffeni albiventer Thomas 

and Wroughton, 1908. See Hayman and 

Hill (1971: 36). 

Holotype. MCZ 8272. Skin and skull. Male. 

Locality. British East Africa ( = Kenya): Meru Riv- 
er, effluent of northern Guaso N^'iro ( = Ewaso Ngi- 
ro). 11 August 1909. 

Collector. G. M. Allen. Original number 113. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Genus PIPISTRELLUS Kaup, 1829 

Eptesicus phasma G. M. Allen, 1911b 

Bull. Mus. Comp. Zool, 54: 327, 

December. 

= Pipistreilus rendalii phasma (G. M. Allen, 

1911). See Koopman (1994: 117). 

Holotype. MCZ 8279. Skin and skull. Male. 

Locality. British East Africa ( = Kenya): Meru Riv- 
er, effluent of northern Guaso Nyiro ( = Ewaso Ngi- 
ro). 6 August 1909. 

Collector. G. M. Allen. Original number 94. 
Condition. Skin and skull complete. 
Type Series. There is a small series of paratypes in 
the MCZ. 

Scabrifer notiusG. M. Allen, 1908 
Bull. Mus. Comp. Zool., 52: 46, July. 
= Pipistreilus capensis notius (G. M. Allen, 
1908). See Koopman (1994: 117). 

Holotype. MCZ 4555. Alcoholic, skull extracted. 
Adult male. 

Locality. South Africa: (Western Cape), Cape 
Town. 

Collector Received from E. L. Lavard, August 
1864. 

Condition. Alcoholic, skull partial (right and left 
zygomatic arches missing; supraoccipital chipped). 
Type Series. Holotyjje only. 

Genus PLECOTUS E. Geoffrey, 1813 

Plecotus sacrimontis G. M. Allen, 1908 
Bull. Mus. Comp. Zool., 52: 50, July 
= Piecotus auritus sacrimontis G. M. Allen, 
1908. See Ognev (1928: 607). 

Holotype. MCZ 6932. Body in alcohol. Adult male. 
Locality. Japan: (Honshu), Mount Fuji. 4 Decem- 
ber 1906.' 



Collector. A. Ovs'ston. 

Condition. Alcoholic. 

Type Series. Holotype only. 

Comments. P. a. sacrimontis was retained as a valid 

subspecies by Koopman (1994: 110). 

Genus SCOTOPHILUS Leach, 1821 

Scotopiiiius altilis G. M. Allen, 1914d 
Bull. Mus. Comp. Zool., 58: 350, July 
= Scotopiiiius leucogaster (Gretzschmar, 
1826). See Koopman (1993: 227). 

Holotype. MCZ 14463. Skin and skull. Adult male. 
Locality. Sudan: Blue Nile, north of (Er) Roseires, 
Aradeiba. 22 January 1913. 

Collector G. M. Allen, Phillips Sudan Expedition. 
Original number 7.3. 
Condition. Skin and skull complete. 
Type Series. 3 paratypes; MCZ 14462, skin and 
skull, male, exchanged to FMNH; and 14610 and 
14611, both males in alcohol. 

Family THYROPTERIDAE Miller, 1907 

Genus THYROPTERA Sp\x, 1823 

Thyroptera tricolor albigula G. M. Allen, 

1923c 

Proc. New England Zool. Club, 9: 1, 10 

December. 

= Thyroptera tricolor albiventer (Tomes, 

1856). See Dunn (1931: 430). 

Holotype. MCZ 20143. Body in alcohol, skull extract- 
ed. Adult female. 

Locality. Panama: Gutierrez, 25 miles (40.2 km) 
inland from Chiriquiscito on trail from Chiriqui La- 
goon, Bocas del Toro to Boquete, Chiriqui. August 
1923. 

Collector E. R. Dunn and C. B. Dvuyea. 
Condition. Alcoholic, skull complete. 
Type Series. 3 paratypes; MCZ 20144, adult male; 
MCZ 20145, juvenile; MCZ 20146, juvenile; all in 
alcohol. 

Order ARTIODACTYLA Owen, 1848 

Family TAYASSUIDAE Palmer, 1897 

Genus PEC/\R/ Reichenbach, 1835 

Tayassu crusnigrum Bangs, 1 902b 
Bull. Mus. Comp. Zool., 39: 20, April. 
^Pecari tajacu crusnigrum (Bangs, 1902). 
See Hershkovitz (1951: 567). 

Holotype. MCZ 10163. Skin and skull. Adult male. 
Locality. Panama: Chiriqui, Boquete ( = Ba)o Bo- 
quete), 4,000 ft (1,220 m). 13 April 1901. 



148 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Collector. W. W. Brown, Jr. Original number 290. 
Condition. Skin and skull complete. 
Type Series. 2 paratypes; MCZ 10162, adult fe- 
male; MCZ 10164, juvenile female. 
Comments. P. t. crusnignim was retained as a valid 
subspecies by Hall (1981: 1080) under the genus 
Dicotijles. 

Tayassu torvus Bangs, 1 898k 

Proc. Biol. Soc. Washington, 12: 164, 10 

August. 

= Pecari tajacu torvus (Bangs, 1898). See 

comments. 

Holotype. B8038. Sldn and skull. Adult male. 

Locality. Colombia: Magdalena, Santa Marta. 26 
January 1898. 

Collector. W. W. Brown, Jr. Original number 50. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Ca- 
brera (1961: 319) under the genus Tayassu. Use of 
the genus Pecari follows Grubb (1993: 380). 

Family MONODONTIDAE Gray, 1821 

Genus DELPHINAPTERUS Lacepede, 
1804 

Beluga declivis Cope, 1865 

Proc. Acad. Nat. Sci. Philadelphia, 17: 

278. 

= Delphinapterus leucas (Pallas, 1776). 

See Hershkovitz (1966: 111). 

Holotype. MCZ 1195. Skull and postcranial skeleton. 
Locality. "Arctic Seas" (probably Greenland). 
Collector. E. K. Kane. 

Condition. Skull partial (right mandibular ramus 
missing). Postcranial skeleton complete except for 
missing left flipper. Right flipper and tail are un- 
cleaned, with tissue largely intact. 
Type Series. Holotype only. 

Comments. Hershkovitz (1966: 111) erroneously 
stated that the holotype of declivis was deposited 
in the Academy of Natural Sciences in Philadel- 
phia, and the holotype of Beluga concreta Cope, 
1865 was in the MCZ. The opposite is in fact true; 
Philadelphia holds the type of B. concreta. 

Family PHOCOENIDAE Gray, 1825 

Genus NEOPHOCAENA Palmer, 1899 

Neomeris asiaeorientalis Pilleri and Gihr, 

1972 

Invest. Cetacea, 4: 126. 

= Neophocaena phocaenoides 

asiaorientalis (Pilleri and Gihr, 1972). See 

van Bree (1973: 17). 

Holotype. MCZ 19998 (but see comments). Skull and 
postcranial skeleton. Adult male. 



Locality. China: Kiangsu (=Jiangsu), Kiangyin, 80 

miles (129 km) northwest of Shanghai. 7 April 

1922. 

Collector F. R. Wulsin. 

Condition. Skin and postcrajiial skeleton complete. 

Type Series. Holotype only. 

Comments. For a discussion of the nomenclature 

and synonymy of this form, consult van Bree 

(1973). Because Neomeris asiaorientalis Pilleri and 

Gihr, 1972 is in fact a replacement name for the 

preoccupied name Delpliinus inelas Schlegel, 1841, 

the holotype of this new name is the same as that 

of Schlegel's name, RMNH 23079. 

Family CERVIDAE Goldfuss, 1820 

Genus ODOCO//.EL/S Rafinesque, 1832 

Cariacus osceola Bangs, 1896b 

Proc. Biol. Soc. Washington, 10: 26, 25 

February. 

= Odocoileus virginianus osceola (Bangs, 

1896). See Lydekker (1915: 148). 

Holotype. B2394. Skin and skull. Adult female. 
Locality. (United States): Florida, Citrus County, 
Citronelle. 29 December 1893. 
Collector. F. L. Small. Original number 1107. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Ttjpe Series. 4 paratypes; B2391, adult male, 
B2392, adult male, B2393, adult female, B2395, 
juvenile male, all represented by skin and skull. 
Comments. O. v. osceola was retained as a valid 
subspecies by Hall (1981: 1096) and Smith (1991: 
1). 

Odocoeleus [sic] virginianus louisianae G. 

M. Allen, 1901 

Amer. Nat., 35: 449. 28 June. 

= Odocoileus virginianus macroura 

(Rafinesque, 1817). See Miller and 

Kellogg (1955: 804). 

Holotype. B9111. Skin and skull. Adult male. 

Locality. (United States): Louisiana, Morehouse 
Parish, Mer Rouge. 8 November 1898. 
Collector B. V. Lilly 

Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; B9112, B8622, B8623, all 
males represented by skin and skull. 

Odocoileus americanus borealis Miller, 

1900 

Bull. New York State Mus. 8: 83, 21 

November. 

= Odocoilus virginianus borealis Miller, 

1900. See Trouessart (1905: 704). 

Holotype. B4999. Skin and skufl. Adult male. 

Locality. (United States): Maine, (Hancock Coun- 
ty), Bucksport. 12 December 1895. 



Type Specimens of Recent Mammals • Helaen and McFadden 



149 



Collector. A. G. Dorr. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. O. v. horealis was retained as a valid 
subspecies bv Hall (1981: 1092) and Smith (1991: 
1). 

Odocoileus virginianus clavium Barbour 

and G. M. Allen, 1922 

J. Mammal., 3: 73, 9 May. 

Holotype. MCZ 19120. Skull and head skin. Adult 
male. 

Locality. (United States): Florida, (Monroe Coun- 
ty), Big Pine Key Winter 1920. 
Collector T. Barbour. 

Condition. Skin and skaill complete. Mandible dis- 
articulated. 

Type Series. 2 paratyjDcs, MCZ 18497, sldn and 
skull, juvenile male; MCZ 18060, sldn and skull, 
juvenile male. 

Comments. Retained as a valid subspecies by Hall 
(1981: 1093) and Smith (1991: 1). 

Genus R/\A/G/FE/=? Hamilton Smith, 1827 

Rangifer arcticus caboti G. M. Allen, 

1914a 

Proc. New England Zool. Club, 4: 104, 

24 March. 

= Rangifer tarandus caribou (Gmelin, 

1788). See Banfield (1962: 70). 

Holotype. MCZ 15372. One shed antler. Adult male. 
Locality. Canada: northeast coast of Labrador, 
about 30 miles (48.3 km) north of Nachvak. 1909. 
Collector O. Biyant. 
Condition. Single antler; complete. 
Type Series. Holotype only. 

Rangifer terraenovae Bangs, 1896! 

Preliminaiy Description of the 

Newfoundland Caribou, Boston, p. 1, 11 

November. 

= Rangifer tarandus caribou (Gmelin, 

1788). See Banfield (1962: 70). 

Holotype. B3778. Skull and head sldn. Adult male. 
Locality. Canada: Newfoundland, Codroy 8 Sep- 
tember 1895 
Collector E. Doane. 

Condition. Skull and head sldn complete. 
Type Series. Bangs remarks that he has "secured a 
series of this fine caribou," corresponding to 
B3779-B3781 and B5757-B5760. 
Comments. J. A. Allen published a description of 
Rangifer terraenovae on 21 November 1896, spec- 
ifying AMNH 11775, a mounted specimen of a 



male adult, as the holotype (1896: 233). Bangs' de- 
scription of this taxon pre-dates J. A. Allen's by 10 
days and thus has priority. 

Family BOVIDAE Gray, 1821 

Genus DAIVIALISCUS Sclater and 
Thomas, 1894 

Damaliscus phillipsi Harper, 1 939 

Proc. Biol. Soc. Washington, 50: 90, 5 

June. 

= Damaliscus pygargus phiillipsi Harper, 

1939. See comments. 

Holotype. MCZ 35443. Skull and skin. Adult male. 
Locality. South Africa: Orange Free State. 23 July 
1935. 

Collector. P. Andreka. Original number 1958d. 
Condition. Skin complete. Skull partial (most of 
palate, left maxilla, and left mandibular ramus 
missing). 

Type Series. Paratype material consists of MCZ 
35444, skin and skull of an adult female as well as 
the following, which Haij)er examined in the col- 
lection of the Academy of Natural Sciences of Phil- 
adelphia; "a mounted head, a skull, and a set of 
horns purchased in Kimberley, Cape Province; a 
mounted head and a set of horns from 'South Af- 
rica'; and two skins and skulls from the Zoological 
Society of Philadelphia." 

Comments. Ansell (1972; 55) used the name Dam- 
aliscus dorcas phillipsi. For the use of pygargns 
over dorcas, see Rookmaaker (1991; 190). 

Order CARNIVORA Bowdich, 1821 

Family CANIDAE Fischer de Waldheim, 
1817 

Genus CANIS Linnaeus, 1758 

Canis lupus beothiucus G. M. Allen and 

Barbour, 1937 

J. Mammal, 18: 230, 14 May. 

Holotype. MCZ 351. Skull and postcranial skeleton. 
Adult, probably male. 

Locality. Canada: Newfoundland. About 1865. 
Collector J. M. Nelson. 

Condition. Skull and postcranial skeleton com- 
plete. 

Type Series. 4 paratypes; 348, skull, adult male; 
349, skull, adult, probably male; 350, skull, adult 
female; MCZ 28726, skin. 

Comments. C. I. beothiicus became extinct around 
1911. Retained as a vahd subspecies by Hall (1981: 
930). 



150 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Pachycyon robustus J. A. Allen, 1885 

Mem. Mus. Comp. Zool., 10: 4, 

December. 

= Canis familiaris Linnaeus, 1758. See G. 

M. Allen (1920a: 498). 

Holotype. MCZ 7091. Postcranial skeleton. 

Locality. (United States): Virginia, Lee County, Ely 
Cave. Probably 1875. 
Collector. N. S. Shaler. 

Condition. Skeleton partial (right scapula, right hu- 
merus, right femur, right tibia, pelvis). 
Type Series. Holotype only. 

Comments. This specimen is a domesticated dog of 
Native Americans. C. lupus familiaris is the name 
now widely used for the domestic dog (Wozencraft 
1993: 281). P. robustus is the type species of Pa- 
chycyon ]. A. Allen, 1885. Pachycyon is a synonym 
oiCanis Linnaeus, 1758, which is commonly over- 
looked, for example, in Wozencraft (199.3) and Mc- 
Kenna and Bell (1997). 

Genus CERDOCYON Hamilton Smith, 
1839 

Cerdocyon thous germanus G. M. Allen, 

1923b 

Proc. Biol. Soc. Washington, 36: 55, 28 

March. 

Holotype. MCZ 19850. Skin and skull. Adult. 

Locality. Colombia: high savannali of Bogota, 9,000 
ft (2,745 m). 

Collector N. Maria. Original number 25. 
Condition. Skin and skull complete. 
Type Series. 5 paratypes; MCZ 19849, skin and 
skull, juvenile; MCZ 20097, sldn and skull, juvenile 
male; 3 specimens from the AMNH are also men- 
tioned in the description. 

Comments. Retained as a valid subspecies by Berta 
(1982: 1). 

Urocyon aquilus Bangs, 1 898h 

Proc. Biol. Soc. Washington, 12: 93, 30 

April. 

= Cerdocyon thous aquilus (Bangs, 1898). 

See Langguth (1969: 178). 

Holotype. B8001. Skin and skull. Adult male. 

Locality. Colombia: (Magdalena), Santa Marta 
Mountains, between 2,000 and 3,000 ft (610-915 
m). 10 February 1898. 

Collector. W. W. Brown, Jr. Original number 58. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B8002, skin and skull, 
adult female. 

Comments. Retained as a valid subspecies by Berta 
(1982: 1). 



Genus L/ROC/OA/ Baird, 1858 

Urocyon cinereoargenteus furvus G. M. 

Allen and Barbour, 1923 

Bull. Mus. Comp. Zool, 65: 266, 

February. 

Holotype. MCZ 19774. Skin and skull. Probably fe- 
male. 

Locality. Panama; Canal Zone, 3 miles (4.8 km) 
west of Balboa. April 1922. 
Collector T. Barbour and W. S. Brooks. 
Condition. Sldn and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Hall 
(1981: 943) and Fritzell and Haroldson (1982; 1). 

Urocyon cinereoargenteus ocyttious 

Bangs, 1899h 

Proc. New England Zool. Club, 1: 43, 5 

June. 

Holotype. B4290. Skin and skull. Adult female. 

Locality. (United States): Wisconsin, Grant Coun- 
ty, Platteville. 25 Januaiy 1896. 
Collector N. E. France. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. Retained as a valid subspecies by Hall 
(1981: 943) and Fritzell and Haroldson (1982: 1). 

Genus UL/LPES Frisch, 1775 

Vulpes deletrix Bangs, 1 898d 

Proc. Biol. Soc. Washington, 12: 36, 24 

March. 

= Vulpes vulpes rubricosa Bangs, 1 898. 

See Churcher (1960: 359). 

Holotype. B6967. Sldn and skull. Adult female. 
Locality. (Canada): Newfoundland, Bay St. 
George. 24 April 1897. 
Collector E. Doane. 
Condition. Sldn and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Vulpes pennsylvanica vafra Bangs, 1 897f 
Proc. Biol. Soc. Washington, 11: 53, 16 
March. 

Name preoccupied by Vulpes vafer 
Leidy, 1869. 



Type Specimens of Recent Mammals • Helpen and McFadden 



151 



Vulpes pennsylvanica rubricosa Bangs, 

1898a 

Science, n. sen, 7: 271, 25 Februaiy. 

(Replacement name for Vulpes 

pennsi/Ivanica vafra Bangs, 1897) 

= Vulpes vulpes rubricosa Bangs, 1898. 

See Churcher (1960: 359). 

Holotype. B116. Skin and skull. Adult female. 

Locality. (Canada): Nova Scotia, Digby. 3 Novem- 
ber 1893. 

Collector. O. Bangs. 
Condition. Skin and skull complete. 
Type Series. 4 paratypes; B1991, skin and skull, and 
B2001, skull, both adult males; B1992, skin and 
sk-ull, and B2002, skull. 

Comments. V. v. rubricosa was retained as a valid 
subspecies by Hall (1981: 939). Vulpes fulvus nib- 
ricatu.s Miller, 1900 (p. 128) was a misspelling and 
thus accidental renaming of Vulpes pennsylvanica 
rubricosa Bangs, 1898. 

Vulpes rubricosa bangsi Merriam, 1900 
Proc. Washington Acad. Sci., 2: 667, 28 
December 28. 

= Vulpes vulpes rubricosa Bangs, 1898. 
See Churcher (1960: 359). 

Holotype. B8880. Skin and skull. Juvenile female. 
Locality. (Canada): Labrador, Lance ( = Lanse) au 
Loup. 2 October 1899. 
Collector E. Doane. 

Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 1 paratype; B8879, skin and skull, 
adult male. 

Family URSIDAE Fischer de Waldheim, 
1817 

Genus L/RSUS Linnaeus, 1758 

Ursus (Euarctos) americanus sornborgeri 

Bangs, 1898j 

Amer. Nat., 32: 500, July. 

= Ursus americanus americanus Pallas, 

1780. See Bangs (1909: 467). 

Holotype. B7411. Skull. Adult, probably female. 
Locality. Canada: Labrador, Okkak ( = Okak). Sum- 
mer 1897. 

Collector J. D. Sornborger, obtained "from the Es- 
kimo." 

Condition. Skull complete. 

Type Series. 2 paratypes; B7412, skull, female; 
B7413, skull, female. 

Comments. A skull from Hopedale, Labrador 
(MCZ 7365), has in the past been erroneously la- 
beled as the holotype of sornborgeri; B7411, the 



true holoty[3e of sornborgeri, is now correctly la- 
beled as such. 

Family PROCYONIDAE Gray, 1825 

Genus PROCYON Storr, 1780 

Procyon gloveralleni Nelson and Goldman, 

1930 

J. Mammal., 11: 453, 11 November. 

= Procyon /otor (Linnaeus, 1758). See 

Corbet and Hill (1991: 104). 

Holotype. MCZ 18591. Skin and skull. Juvenile male. 
Locality. Barbados. 1920. 
Collector F. Watts. 
Condition. Skin and skull complete 
Type Series. Holotype only. 

Comments. Considered a valid species by Wozen- 
craft (1993: 335) and Nowak (1999: 698) but almost 
certainly introduced to Barbados in the 17th cen- 
tuiy (Helgen and Wilson, in prep.). The last rac- 
coon on Barbados was seen in 1964, and the pop- 
ulation is probably extinct. 

Procyon lotor elucus Bangs, 1898b 
Proc. Boston Soc. Nat. Hist., 28: 219, 15 
March. 

Holotype. B3502. Skin and skull. Adult male. 

Localitij. (United States): Florida, Brevard County, 
Oak Lodge, east peninsula opposite Micco. 15 Feb- 
ruary 1895. 
Collector. O. Bangs. 

Condition. Skin complete. Skull partial (condyle, 
coronoid, and angular processes of left mandibular 
ramus broken). Mandible disarticulated. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. Retained as a valid subspecies by Hall 
(1981: 968). 

Procyon maynardi Bangs, 1898g 

Proc. Biol. Soc. Washington, 12: 92, 30 

April. 

= Procyon /otor(Linneaus, 1758). See 

Koopman et al. (1957: 164). 

Holotype. B7750. Skin and skull. Juvenile male. 
Locality. Bahamas: New Providence Island, Nas- 
sau. August 1897. 
Collector H. L. Claridge. 

Condition. Skin complete. Skull partial (broken 
from frontals to occiput). Mandible disarticulated. 
Type Series. Holotype only. 

Comments. Considered a valid species by Wozen- 
craft (1993: 336) and Nowak (1999: 698) but un- 
doubtedly a recent introduction to New Providence 
Island (see Olson and Pregill, 1982: 5). 



152 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Family MUSTELIDAE Fischer de 
Waldheim, 1817 

Genus LONTRA Gray, 1843 

Lutra degener Bangs, 1 898d 

Proc. Biol. Soc. Washington, 12: 35, 24 

March. 

= Lontra canadensis canadensis 

(Sclireber, 1776). See van Zyll de Long 

(1972: 81). 

Holotype. B6965. Sldn and skull. Adult male. 

Locality. Canada: Newfoundland, Bay St. George. 

23 April 1897. 

Collector. E. Doane. 

Condition. Skin and skull complete. 

Type Series. Paratype material consists of B6966, 

skin and skull of an adult female, mentioned by 

number in the original description, as well as "two 

extra skulls," corresponding to B3755 and B3799, 

and "a large series of unsexed otter skulls from 

Newfoundland," corresponding to MCZ 494—508. 

Lutra hudsonica vaga Bangs, 1 898b 

Proc. Boston Soc. Nat. Hist., 28: 224, 15 

March. 

= Lontra canadensis laxatina F. Cuvier, 

1823. See van Zyll de Long (1972: 81). 

Holotype. B5749. Skin and skull. Adult male. 

Locality. (United States), Florida, Brevard Count)-, 

Micco. 17 March 1897. 

Collector F. R. Hunter. 

Condition. Skin and skull complete. 

Type Series. 4 paratypes; B4995, skin and skull, 

adult female; B4998, sldn and skull, adult male; 

B6092, skin and skull, adult male; B6093, sldn and 

skull, adult female. 

Genus MARTES Pinel, 1792 

Mustela atrata Bangs, 1897b 

Amer. Nat., 31: 162, 1 February. 

= Martes americana atrata (Bangs, 1897). 

See G. M. Allen (1942: 166) 

Holotype. B5752. Skin and skull. Adult female. 

Locality. Canada: Newfoundland, Bay St. George. 
29 September 1896. 

Collector E. Doane. Original number 2. 
Condition. Sldn and skull complete. 
Type Series. 12 paratypes; B5751, skin and skull, 
adult female; MCZ 492-93, 509-517, unsexed 
skulls. 

Comments. M. a. atrata was retained as a valid sub- 
species by Hall (1981: 983). 



Mustela brumalis Bangs, 1898j 

Amer. Nat., 32: 502, July 

= Martes americana atrata (Bangs, 1897). 

See Clark et al. (1987: 1). 

Holotype. B7417. Skull. Adult, probably male. 

Locality. Canada: Labrador, Okkak ( = Okak). Sum- 
mer 1897. 

Collector J. D. Sornborger, obtained "from the Es- 
kimo." 

Condition. Skull complete. 

Type Series. 2 paratypes; B7418, skull; B7419, 
skull; both probably male. 

Genus MUSTELA Linnaeus, 1758 

Mustela cicognanii mortigena Bangs, 1913 
Bull. Mus. Comp. ZooL, 54: 511, July 
= Mustela erminea rictiardsonii Bonaparte, 
1838. See Hall (1951: 110). 

Holotype. B3745. Skin and skull. Adult male. 

Locality. Canada: Newfoundland, Bay St. George. 

27 September 1895. 

Collector. E. Doane. Original number 1. 

Condition. Skin and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Putorius frenatus neomexicanus Barber 

and Cockerell, 1898 

Proc. Acad. Nat. Sci. Philadelphia, 1898, 

p. 188, May 

= Mustela frenata neomexicana (Barber 

and Gocl<erell, 1898). See Miller (1912: 

100). 

Holotype. MCZ 10475. Skin with extra tail, and skull. 
Adult male. 

Locality. (United States): New Mexico, (Dona Ana 
County), Mesilla, shore of Armstrongs' Lake, 3,800 
ft (1,159 m). 1 Februaiy 1898. 
Collector A. C. Tyson. Original number 58. 
Condition. Sldn complete, but tail poorly prepared. 
The tail belonging to a discarded topotype is tied 
to the holotype as an example. Skull complete. 
Type Series. A topotype taken at the same time as 
the holotype was partially decomposed and dis- 
carded; the tail of this specimen is included with 
MCZ 10475. The original description also refers to 
"a specimen, without any histoiy, in alcohol ... in 
the collection of the New Mexico Agricultural Col- 
lege." 

Comments. M. f. neomexicana was retained as a 
valid subspecies by Hall (1981: 995). 



Type Specimens of Recent Mammals • Helpen and McFadden 153 



Putorius (Arctogale) longicauda oribasus 

Bangs, 1899m 

Proc. New England Zool. Club, 1: 81, 27 

December. 

= Mustela frenata oribasus (Bangs, 1899). 

See Hall (1936: 105). 

Holotype. B9058. Skin and skull. Adult female. 

Locality. Canada: British Cokmibia, source of Ket- 
tle River, 7,500 ft (2,288 m). 10 September 1898. 
Collector. A. C. Brooks. Original number 1368. 
Conclition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. M. f. oribasus was retained as a valid 
subspecies by Hall (1981: 998). 

Putorius (Lutreola) lutensis Bangs, 1 898b 

Proc. Boston Soc. Nat. Hist., 28: 229, 15 

March. 

= Musteia vison lutensis (Bangs, 1898). 

See Hollister (1913: 474). 

Holotype. B7225. Skin and skull. Adult male. 

Localitij. (United States), Florida, St. Johns Coun- 
ty, salt marsh opposite Matanzas Inlet. 16 February 
1897. 

Collector. O. Bangs. Original number 7. 
Condition. Skin and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. M. v. lutensis was retained as a valid 
subspecies by Hall (1981: 1003) and Lariviere 
(1999: 1). 

Putorius (Arctogale) muricus Bangs, 1 899j 

Proc. New England Zool. Club, 1: 71, 31 

July. 

=Mustela erminea muricus (Bangs, 1899). 

See Hall (1945: 77). 

Holotype. B9146. Skin and skull. Juvenile male. 
Locality. (United States): California, El Dorado 
County, Echo, 7,500 ft (2,288 m). 15 July 1897. 
Collector W. W. Price and E. M. Nutting. Original 
number 266. 

Conclition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. Holotype only. 

Comments. M. e. muricus was retained as a valid 
subspecies by Hall (1981: 990) and King (1983: 1). 

Putorius noveboracensis notius Bangs, 

1899i 

Proc. New England Zool. Club, 1: 53, 9 

June. 

= Mustela frenata noveboracensis 

(Emmons, 1840). See Hall (1936: 104). 

Holotype. B2678. Skin and skull. Juvenile male. 
Locality. (United States): North Carolina, Bun- 
combe County, Weavei"ville. 10 July 1892. 



Collector. J. S. Cairns. Original number 2214. 
Condition. Sldn complete. Skull partial (two frag- 
ments only, premaxilla— lachrymal). Mandible dis- 
articulated. 

Type Series. 2 paratypes; AMNH 1247, adult male; 
USNM 32239, adult male. 

Putorius occisor Bangs, 18991 

Proc. New England Zool. Club, 1: 54, 9 

June. 

= Mustela frenata occisor {Bangs, 1899). 

See Hall (1936: 104). 

Holotype. B9102. Skin and skull. Adult male. 

Locality. (United States): Maine, Hancock County, 
Bucksport, near mouth of Penobscot River. 15 Jan- 
uary 1899. 

Collector A. G. Dorr. 

Condition. Skin complete. Skull slightly damaged 
(left zygomatic arch broken). 

Type Series. There is a series of paratypes in the 
MCZ. 

Comments. M. f. occisor was retained as a valid 
subspecies by Hall (1981: 997). 

Putorius rixosus Bangs, 1896a 

Proc. Biol. Soc. Washington, 10: 21, 25 

Februaiy. 

= Mustela nivalis rixosa (Bangs, 1896). 

See Reichstein (1958: 169). 

Holotype. B642. Skin and skull. Adult female. 

Locality. Canada: Saskatchewan, Osier 15 July 
1893. 

Collector W. C. Colt. Original number 79/181. 
Condition. Skin complete. Skull slightly damaged 
(left zygomatic arch broken). 

Type Series. Three specimens other than the ho- 
lotype are mentioned by number in the original 
description; MCZ 5532, USNM 4231, probably fe- 
male, and USNM 13904, probably male. All are 
unsexed skins. 

Comments. M. n. rixosa was retained as a valid sub- 
species by Hall (1981: 993) and Sheffield and King 
(1994: 1). 

Putorius vison energumenos Bangs, 

1896c 

Proc. Boston Soc. Nat. Hist., 27: 5, 

March. 

= Mustela vison energumenos (Bangs, 

1896). See Miller (1912: 101). 

Holotype. B3555. Skin and skull. Adult male. 

Locality. Canada, British Columbia, Sumas. 23 
September 1895. 

Collector A. C. Brooks. Original number 514. 
Condition. Skin and skull complete. 



154 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Type Series. 1 paratype; B3556, skin and skull, ju- 
venile male. 

Comments. M. v. energumenos was retained as a 
valid subspecies by Hall (1981: 1001) and Lariviere 
(1999: 1). 

Putorius (Lutreola) vulgivagus Bangs, 

1895b 

Proc. Boston Soc. Nat. Hist., 26: 539, 31 

July. 

= Mustela vison vulgivaga (Bangs, 1895). 

See Miller (1912: 102). 

Holotype. B2751. Skin and skull. Adult male. 

Locality. (United States): Louisiana, Plaquemines 
Parish, Burbridge. 10 Januaiy 1895. 
Collector. F. L. Small. Original number 1439/54. 
Condition. Skin and skull complete. 
Tijpe Series. 10 paratypes; B2752-B2761, all rep- 
resented by sldn and skull, 9 males and 1 female. 
Comments. M. v. vulgivaga was retained as a valid 
subspecies by Hall (1981: 1004) and Lari\'iere 
(1999: 1). 

Putorius xanthogenys mundus Bangs, 

1899i 

Proc. New England Zool. Club, 1: 56, 9 

June. 

= Mustela frenata munda (Bangs, 1899). 

See Hall (1936: 107). 

Holotype. B5459. Skin and skull. Adult male. 

Locality. (United States): California, Marin Coun- 
ty, Point Reyes. 19 June 1896. 
Collector C. A. Allen. Original number 931. 
Condition. Skin complete. Skull slightly damaged 
(left zygomatic arch broken). 

Type Series. 1 paratype, B8632 (not B8631, men- 
tioned erroneously in the original description), skin 
and skull, male. 

Comments. M. f. munda was retained as a valid 
subspecies by Hall (1981: 995). 

Family MEPHITIDAE Bonaparte, 1845 

Genus MEPHITIS E. Geoffrey and G. 
Cuvier, 1795 

Mephistis avia Bangs, 1898c 

Proc. Biol. Soc. Washington, 12: 32, 24 

March. 

=- Mephitis mephitis avia Bangs, 1898. See 

Hall (1936: 65). 

Holotype. B5747. Skin and skull. Adult male. 

Locality. (United States): Illinois, Mason County, 
San Jose. 10 March 1897. 

Collector. H. H. and C. S. Brimley Original num- 
ber 2500. 



Condition. Skin and skull complete. 
Ti/pe Series. 1 paratype; B5783, skin and skull, 
adult male. 

Comments. M. m. avia was retained as a valid sub- 
species by Hall (1981: 1019) and Wade-Smith and 
Verts (1982: 1). 

Mephitis mephitica elongata Bangs, 1895b 

Proc. Boston Soc. Nat. Hist., 26: 531, 31 

July. 

= Mephitis mephitis elongata Bangs, 1895. 

See A. H. Howell (1921: 39). 

Holotype. B3051. Skin and skull. Adult male. 

Locality. (United States): Florida, Brevard County, 

Micco. 5 March 1895. 

Collector O. Bangs. 

Condition. Skin and skull complete. 

Type Series. There is a series of paratypes in the 

MCZ. 

Comments. Retained as a valid subspecies by Hall 

(1981: 1019) and Wade-Smith and Verts (1982: 1). 

Mephitis mephitica scrutator Bangs, 1 896i 

Proc. Biol. Soc. Washington, 10: 141, 28 

December. 

= Mephitis mephitis mesomelas 

Lichtenstein, 1832. See Hall (1936: 66). 

Holotype. B2889. Skin and skull. Adult male. 

Locality. (United States): Louisiana, Acadia Parish, 
Caitville. 25 May 1895. 

Collector F. L. Small. Original number 1842. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Ti/pe Series. 1 parats-pe, B2886, skin and skull, 
adult female. 

Mephitis spissigrada Bangs, 1898c 

Proc. Biol. Soc. Washington, 12: 31, 24 

March. 

= Mephitis mephitis spissigrada Bangs, 

1898. See Hall (1936: 67). 

Holotype. B3699. Skin and skull. Adult female. 

Locality. Canada: British Columbia, Sumas. 30 
September 1895. 

Collector A. C. Brooks. Original number 518. 
Condition. Skin and skull complete. Mandible dis- 
articulated. 

Type Series. 3 paratypes; B3700, skin and skull, 
adult female, and B5548, skin and skull, adult male; 
B7435, skull, adult male. 

Comments. M. m. spissigrada was retained as a val- 
id subspecies by Hall (1981: 1022) and Wade- 
Smith and Verts (1982: 1). 



Type Specimens of Recent Mammals • Hel^en and McFadden 155 



Genus SPILOGALE Gray, 1865 

Spilogale ambarvalis Bangs, 1 898b 

Proc. Boston Soc. Nat. Hist., 28: 222, 15 

March. 

= Spilogale putorius ambarvalis Bangs, 

1898. See Van Gelder (1953: 255). 

Holotype. B3481. Skin and skull. Adult male. 

Locality. (United States): Florida, Brevard County, 

Oak Lodge, east peninsula opposite Mieco. 30 Jan- 

uaiy 1895. 

Collector. O. Bangs. Original number 11. 

Condition. Skin and skull complete. 

Type Series. There is a large series of paratypes in 

the MCZ. 

Comments. S. p. ambarvalis was retained as a valid 

subspecies by Hall (1981: 1014) and Kinlaw (1995: 

1). 

Family VIVERRIDAE Gray, 1821 

Genus EUPLERES Doyere, 1835 

Eupleres major Layau6en, 1929 

Compt. Rend. Acad. Sci. Paris, 189: 198, 

22 July 22. 

= Eupleres goudotii major Lavauden, 

1929. See Albignac (1973: 23). 

Syntypes. MCZ 45691: Skin, skull, and postcranial 
skeleton. Subadult female, MCZ 45962: Skin, skull, 
and postcranial skeleton. Subadult male. 
Locality. Madagascar: (Antsiranana), foot of the 
Massif Tsaratanna (=Tsaratanana), Upper Sombi- 
rano Valley, above village of Beangona, 1,500 m. 
April 1929. 
Collector Lavauden. 

Condition. MCZ 45691: Skin complete, v^dth bald 
spot on dorsum and tail slightly damaged. Skull and 
skeleton complete. Mandible disarticulated. Teeth 
removed from skull but present. MCZ 45962: Skin 
complete. Skull and skeleton complete. Mandible 
disarticulated. Teeth removed from skull but pres- 
ent. 

Type Series. 2 syntypes, described above. 
Comments. These are the two specimens from G. 
Grandidier's personal collection on which Lavau- 
den based his original description of Enpleres ma- 
jor Albignac (1973: 23) wrote that these type spec- 
imens were "introuvable [nowhere to be found]." 

Family HERPESTIDAE Bonaparte, 1845 

Genus GALIDICTIS I. Geoffrey, 1839 

Galidictis grandidiensis [sic] Wozencraft, 

1986 

J. Mammal. 67: 561, 8 August. 

= Galidictis grandidieri Wozencraft, 1 986. 

See Wozencraft (1987: 198). 

Holotype. MCZ 45983. Skin, skull, and postcranial 
skeleton. Adult. 



Locality. Madagascar (no further data available). 
The locahty of the paratype, stored in the AMNH, 
is "Madagascar, Lac Tsimanampetsotsa, 24°08' S, 
43°46' E." 

Collector No collection data available. The holo- 
type is part of the collection of G. Grandidier. 
Condition. Skin prepared flat; incomplete (ven- 
trum missing). Skull complete. 
Type Series. 1 paraty^^e; AMNH 100478, sldn and 
skull, adult male. 

Comments. G. grandidieri was considered a valid 
species by Wozencraft (1993: 300) and Nowak 
(1999: 769). 

Family FELIDAE Fischer de Waldheim, 

1817 

Genus LEPTAILURUS Sevenzoy, 1858 

Fells capensis phillipsi G. M. Allen, 1914d 

Bull. Mus. Comp. Zool., 58: 337, July. 

= Leptallurus serval phillipsi (G. M." Allen, 

1914). See comments. 

Holotype. MCZ 14908. Skin and skeleton. Adult 
male. 

Locality. Sudan: Blue Nile, El Garef 10 Januaiy 
1913. 

Collector J. C. PhiUips. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. G. M. Allen (1939: 241) used the name 
Felis serval phillipsi. The use of the genus Lep- 
tailunis follows Wozencraft (1993: 292). L. s. phil- 
lipsi was retained as a valid subspecies by Smithers 
(1975: 7). 

Genus LVA/XKerr, 1792 

Lynx (Cervaria) fasciatus oculeus Bangs, 

1899e 

Proc. New England Zool. Club, 1: 23, 31 

March. 

= Lynx rufus callfornlcus Mearns, 1897. 

See Grinnell and DLxon (1924: 346). 

Holotype. B8633. Skin and skull. Adult male. 

Locality. (United States): Cahfomia, Marin Coun- 
ty, Nicasio. 11 December 1898. 
Collector C. A. Allen. Original number 981. 
Condition. Skin and skull complete. 
Type Series. 1 paratype; B4789, sldn and skull, 
adult male. 

Lynx gigas Bangs, 1897e 

Proc. Biol. Soc. Washington, 11: 50, 16 

March. 

= Lynx rufus gigas Bangs, 1897. See 

Peterson and Downing (1952: 11). 

Holotype. B4951. Skin and skull. Adult male. 

Locality. Canada: Nova Scotia, 15 miles (24.1 km) 
back of Bear River. 11 December 1895. 



156 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Collector. D. R. Ritchie. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. L. r gigas was retained as a valid sub- 
species by Hall (1981: 1054). 

Lynx subsolanus Bangs, 1897e 

Proc. Biol. Soc. Washington, 11: 49, 16 

March. 

= Lynx canadensis subsolanus Bangs, 

1897. See ElHot (1901: 296). 

Holotype. B1190. Sldn and skull. Adult male. 

Locality. Canada: Newfoundland, Codroy. 13 June 
1894. 

Collector E. Doane. 
Condition. Skin and skull complete. 
Type Series. 2 paratypes; B5754, skin and skull, ju- 
venile female; B3798, skull, adult male. 
Comments. L. c. subsolanus was retained as a valid 
subspecies by Hall (1981: 1051). 

Genus PUMA Jardine, 1834 

Felis bangs! Merriam, 1901 

Proc. Washington Acad. Sci., 3: 595, 11 

December. 

= Puma concolor bangs! {Memam, 1901). 

See coinments. 

Holotype. B8413. Skin and skull. Adult male. 

Locality. Colombia: (La Guajira), Dibulla. 8 Oc- 
tober 1899. 

Collector. W. W. Brown, Jr. 
Condition. Skin and skull complete. 
Type Series. 3 paratypes; B8147, skin and skull, 
adult female, and "two skulls from Peru, in the 
American Museum of Natural History." 
Comments. Nelson and Goldman (1929: 347) used 
the name Felis concolor bangsi. The use of the ge- 
nus Puma follows Wozencraft (1993: 296). Re- 
tained as a valid subspecies by Currier (1983: 1). 

Fells bangsi costaricensis Merriam, 1901 

Proc. Washington Acad. Sci., 3: 596, 11 

December. 

= Puma concolor costaricensis (Merriam, 

1901). See comments. 

Holotype. MCZ 10118. Skin and skull. Adult female. 
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo- 
quete), 4,000 ft (1,220 m). 22 April 1901. 
Collector W. W. Brown, Jr. Original number 337. 
Condition. Skin and skull complete. 
Type Series. There is a series of paratypes in the 
MCZ. 

Comments. Nelson and Goldman (1929: 347) used 
the name Felis concolor costaricensis. The use of 
the genus Puma follows Wozencraft (1993: 296). 
Retained as a valid subspecies by Currier (1983: 1). 



Felis coryi Bangs, 1899b 

Proc. Biol. Soc. Washington, 13: 15, 31 

Januaiy. 

= Puma concolor coryi {Bangs, 1899). See 

comments. 

Holotype. B7742. Sldn and skull. Adult male. 

Locality. (United States): Florida, Brevard County, 
"wildeniess back of Sebastian". 1 Januaiy 1898 
Collector F. R. Hunter. 
Condition. Skin and skull complete. 
Type Series. 5 paratypes; B5489, adult female, 
B5650, adult female, B6992, aduh male, B7743, 
adult female, B7744, juvenile female; all repre- 
sented by sldn and skull. 

Comments. Nelson and Goldman (1929: 347) used 
the naine Felis concolor coryi; the use of the genus 
Puma follows Wozencraft (1993: 296). Felis coryi 
Bangs, 1899 is a replacement name for Felis con- 
color floridana Cory, 1896 (1896: 109). Retained as 
a valid subspecies by Currier (1983: 1). 

Felis improcera Phillips, 1912 

Proc. Biol. Soc. Washington, 25: 85, 4 

May. 

= Puma concolor improcera (Phillips, 

1912). See comments. 

Holotype. MCZ 12704. Skin and skull. Adult male. 
Locality. (Mexico): Lower ( = Baja) California, Cal- 
malli. 3 September 1911. 

Collector. E. W Funcke. Original number 10. 
Condition. Skin and skull complete. 
Type Series. Holotype only. 

Comments. Nelson and Goldman (1929: 347) used 
the name Felis concolor improcera. The use of the 
genus Puma follows Wozencraft (1993: 296). Re- 
tained as a valid subspecies by Currier (1983: 1). 

Order CIMOLESTA McKenna, 1975 

Family MANIDAE Gray, 1873 

Genus PH/Ar/\G/A/aS Rafinesque, 1821 

Phataginus tricuspis mabirae G. M. Allen 

and Loveridge, 1942 

Bull. Mus. Comp. Zool, 89: 178, 

Februaiy. 

Holotype. MCZ 39417. Skin, skull, and postcranial 
skeleton. Adult male. 

Locality. Uganda: (Buganda), Chagwe, Mabira 
Forest, Mubango. 12 November 1938. 
Collector A. Loveridge. 

Condition. Skin and skull complete. Postcranial 
skeleton partial (includes atlas, right tibia, and right 
fibula). 
Type Series. Holotype only. 



Type Specimens of Recent Mammals • Helgen and McFadden 157 



Comments. Retained as a valid subspecies by 
Meester (1972: 2). 

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INDEX 



Type Specimens of Recent Mammals • Hel^en and McFadden 171 



abaconis 

Geocapromijs, 132 
abbotti 

Mijotis, 146 
abbreviata 

Neotonia, 121 
aberti 

Sciunis, 99 
abietorum 

Feromyscus, 125 
ablutus 

Leggada, 118 
acadicus 

Microtus, 115 

Neosorex, 138 

Zapus, 112 
aequalis 

Aedialops, 140 

Scalops, 140 

Scalopus, 140 
aestuans 

GueHinguetus, 106 

Sciiirus, 106 
Aethalops 

aequalis, 140 

alecto, 140 
Aethoglis 

argenteus, 129 

/jueij, 129 
Aethosciurus 

byatti, 106 

laetus, 106 
africanus 

Nycticeius, 147 
Afrosoricida, 101 
AgOMfi 

paca, 130 

virgatus, 130 
Agoutidae, 130 
akka 

FuniscUinis, 104 
Akodon 

apricus, 127 

teguina, 127 

xerampelinus, 127 
alacer 

Lepus, 134 

Sylvilagus, 134 
alashanicus 

Spermophilus, 107 
albicinctus 

Myotis, 146 
albigula 

Thyroptera, 147 
albiventer 

Nycticeius, 147 



Thyroptera, 147 
alboniger 

Hylopetes, 105 

Fteroinys, 105 
alecto 

Aethalops, 140 
alfari 

Microsciunis, 105 
alleni 

Dasymys, 118 

Heteromys, 112 

Hylomyscus, 118 

Lepus, 133 

Liomys, 112 

Macrotolagus , 133 

Melomys, 118 

Rhinolophus, 143 

rt/ogfl 

Blarina, 137 
AZoKflrtfl 

luctuosa, 135 

palliata, 135 

pigra, 135 

trabeata, 135 
alpinus 

Sciuroptenis, 104 
altilis 

Scotophilus, 147 
amazonicus 

Nectomys, 121 
ambarvalis 

Spilogale, 155 
americana 

Martes, 152 
americanus 

Euarctos, 151 

Lept/s, 133, 134 

Odocoileus, 148 

Ursus, 151 
A77iefn(ia 

centurio, 143 

minor, 143 
ammodijtes 

Feromyscus, 125 
anastasae 

Feromyscus, 124, 125 

Scalops, 140 

Scalopus, 140 
anchietae 

Otomys, 120 
anchises 

Hylopetes, 105 

Fteromys, 105 
anetianus 

Fteropus, 141 
angolensis 

Mops, 145 



Antechinus 

mayeri, 100 

misim, 100 

noso, 100 
Antillogale 

marcanoi, 137 
aorensis 

Fteropus, 141 
Aplodontia, 103 

califomica, 103 

Columbiana, 103 

rainieri, 103 

7^(/a, 103 
Aplodontidae, 103 
Apodemus 

euxinus, 117 

mystacinus, 117 
apricus 

Akodon, 127 

Scotinomys, 127 
aquaticus 

Scalopus, 140 
aquilis 

Cerdocyon, 150 

Craseomys, 113 

Urocyon, 150 
aquilonius 

Fiber, 116 

Ondatra, 116 
araeum 

Flagiodonta, 132 
araneus 

Sorex, 138 
arcticus 

Hesperomys, 124 

Lepus, 133 

Rangifer, 149 
ArctogaZe 

longicauda, 153 

muricus, 153 

oribasus, 153 
Arcto??ii/.s 

avanis, 105 

flaviventer, 105 

igna-uus, 105 
arenarius 

Feromyscus, 126 
argentatus 

Feromyscus, 125 
argenteus 

Aethoglis, 129 

Graphiurus, 129 
arJeZ 

Fteropus, 141 
armatus 

Spermophilus, 107 
A/tiZjeus 



femurvillosum, 144 

lituratus, 144 

palmarum, 144 
Aitiodactyla, 145 
Arujco/a 

breweri, 114 

longipilis, 114 

riparia, 114 

rufidorsum, 114 

terraenovae, 114 
Arvicolinae, 112 
aslaeorientalis 

Neomeris, 148 

Neophocaena, 148 
Atalapha 

brachyotis, 146 

Suncus, 139 
atrata 

Martes, 152 

Mustela, 152 
atrodorsalis 

Thomomys, 111 
attioateri 

Feromyscus, 124 

Geogale, 101 
Triaenops, 143 
auritus 

Flecotus, 147 

Eothenomys, 113 

Microtus, 113 
austerulus 

Sigmodon, 127 
austini 

Fteropus, 141 
australis 

Cryptogale, 101 

Echymipera, 101 

Reithrodontomys, 127 
austrinus 

Geomys, 110 

Mannota, 105 
Arctomys, 105 

Cryptotis, 138 

Mephitis, 154 
bactrianus 

Mus, 121 
hadlus 

Ictidonujs, 108 

Spennophilus, 108 
hairdii 

Lepus, 133, 134 



172 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Mus, 98 

Peromyscii.s, 98 
haliolus 

Peromyscus, 126 
baluensis, 

Callosciunis, 103 
bangsi 

Fells, 156 

Glaucomys, 104 

Perognathus, 112 

Puma, 156 

Sciuroptenis, 104 

Viilpes, 151 
bangsii 

Lepus, 133 
barbouri 

Otomys, 121 
barrerae 

Octoniys, 130 

Tympanoctomys, 130 
Bathyergidae, 129 
hella 

Neotoma, 121 
belliis 

Peromyscus, 124 
Beluga 

concreta, 148 

declivis, 148 
beothucus 

Cants, 149 
bicolor 

Crocidura, 137 

Hipposideros, 142 

Oecomys, 122 
Blarina 

aloga, 137 

brevicauda, 137 

compacta, 137 
borealis 

hasiurus, 146 

Odocoileus, 148 

Peromyscus, 124 

Synaptomys, 116 

Tamias, 109 
Boromys 

torrei, 131 
borucae 

Sigmodon, 127 
Bovidae, 149 
brachyotis 

Atalapha, 146 

Lasiurus, 146 
brasiliensis 

Sylvilagus, 134 
brevicauda 

Blarina, 137 

Zygodontomys, 128 
brevicaudata 

Microgale, 102 

Arvicola, 114 



Microtus, 114 
brochus 

Syntheosciunis, 108 
browni 

Microsciunis, 105 

Sciurus, 105 
bninialis 

Mai-tes, 152 

Mustela, 152 
biifo 

Leggada, 118 

Mus, 118 
burrus 

Proechimys, 131 
byatti 

Aethosciurus, 106 

Paraxerus, 106 
caboti 

Rangifer, 149 
cacabatus 

Peromyscus, 124 

Castor, 109 
caecutiens 

Sorex, 138 
cfi/er 

Pedetes, 128 
californica 

Aplodontia, 103 
califoiiticus 

Dipodomys, 111 

Lynx, 165 

Scapanus, 140 
callida 

Dasyprocta, 130 
Callosciunis 

baluensis, 103 

erythraeus, 103 

ferrugineus, 103 

haemobaphes, 103 

medialis, 103 

primus, 103 
Caluromyidae, 99 
Caluromys 

cicur, 99 

lanatus, 99 
canadensis 

Castor 109 

Lontra, 152 

Lynx, 156 

Peromyscus, 125 
Canidae, 149 
canina 

Peropteryx, 142 
Cfinzs 

beothucus, 149 

familiaris, 150 

/tipus, 149, 150 
Cansumys 

canus, 117 
canus 



Cansumys, 117 
capensis 

Fells, 165 

Pedetes, 128 

Pipistrellus , 146 
Capromyidae, 131 
Capromys 

nana, 132 

pilorides, 131 

relictus, 131 
capucinus 

Cebus, 136 
caraccioli 

Vampyrodes, 145 
Cariacus 

osceola, 148 
caribou 

Rangifer, 149 
carissinia 

My Otis, 146 
Carnivora, 149 
carolinensis 

Sciurus, 106 
carpenteri 

Hijlobates, 136 
carroruni 

Oryzomys, 123 
cascadensis 

Lepus, 134 
castaneoventris 

Sciurus, 103 
castanonotus 

Sciurus, 99 
castanotus 

Sciurus, 99 
casta nops 

Cratogeomys, 111 

Pappogeomys, 111 
Castor 

caecator, 109 

canadensis, 109 
Castoridae, 109 
cavator 

Macrogeomys, 110 

Orthogeomys, 110 
cayennensis 

Proechimys, 131 
Cebidae, 135 
Cebus 

capucinus, 136 

curtus, 136 
celatus 

Phenacomys, 116 
centu rlo 

Ametrida, 143 
Cerdocyon 

aquilus, 150 

germanus, 150 

thous, 150 
Cervaria 

fasclatus, 155 



oculeus, 155 
Cei-vidae, 148 
Cetartiodactyla, 97 
Chaerophon 

leucostigma, 145 
Cheiromys 

laniger, 135 

madagascariensls, 135 
c/ierriei 

Zygodontomys, 128 
Chilonycteris 

pamellii, 143 

pusillus, 143 

torrei, 143 
Chionomys 

nivalis, 112 
chionopaes 

Dicrostonyx, 113 
chiriquensis 

Guerlinguetus, 106 

Sclunis, 106 
Chiroptera, 140 
Chlorotalpa 

tropicalis, 101 
chrotorrhinus 

Microtus, 114 
Chrysochloridae, 101 
Chnjsochloris 

stidilmanni, 101 

tropicalis, 101 
chrysogaster 

Lenimus, 114 
cicognonn 

Mustela, 152 
cicur 

Caluromys, 99 

Philander, 99 
Cimolesta, 156 
cinereoargenteus 

Urocyon, 150 
cJnereHS 

Sorgx, 139 
Cingulata, 101 
Citellus 

obscurus, 107 

.siccus, 107 
Clavlglls 

collaris, 129 

soleatus, 129 
clavlum 

Odocoileus, 149 
Clethrionomys 

gapperi, 113 

proteus, 113 
collaris 

Claviglis, 129 
colondylana 

Dasyprocta, 130 
colonus 

Geomys, 110 
coloratus 



Type Specimens of Recent Mammals • Hclfien and McFadden 173 



On/zoini/s, 123 
coliuiibiaiia 

Aplodoiifia, 103 
coliimbicnnis 

Geocapromijs, 132 
compacta 

Blarina, 137 
concoJor 

Felis, 156 

Puma, 156 
concreta 

Beluga, 148 
condylunts 

Mops, 145 
confucianus 

Niviventer, 119 
consobrinus 

Sciurofamias, 106 
cooper; 

Synaptovujs, 116 

Tainias, 109 
cory; 

Fe/;.s, 156 

Puma, 156 
Conjnorhinus 

phijllotis, 146 
casta ricensis 

Felis, 156 

Pinna, 156 
cowani 

Microgale, 102 
Craseomijs 

aquilus, 113 
crassus 

Phenacornys, 116 
Cratogeomys 

castanops. 111 

rubellus. 111 
crgper 

ReUhrodontomys , 127 
Cricetinae, 117 
Cricetomylnae, 117 
cricefulus 

Saccostomus, 117 
crinitus 

Peromyscus, 125 
Crocidura 

bicolor, 137 

fuscomurina, 137 

geata, 138 

hildegardeae, 138 

maurisca, 138 

phaios, 138 

tephragaster, 137 
cnisiiignim 

Pecan, 147 

Tayassu, 147 
Cryptogale 

a u straits, 101 
Cryptomys 

hottentotus, 129 



occlusus, 129 

ivhijtei, 129 
Cryptotis 

avia, 138 

tJioinasi, 138 
cubanus 

Geocaproinys, 132 

Solenodon, 137 
cumbedandius 

Geoim/s, 110 
c;/ppra 

Ochotona, 133 
ctirttis 

CeZpjf.s, 136 

Hipposideros, 142 
Damaliscus 

pJiiUipsi, 149 

pygargus, 149 
daphaenodon 

Sorex, 139 
darlingtoni 

Eptesicus, 145 

PipistreUus, 145 
Dasogale 

fontoynonti, 103 
Dasyinys 

alleni, 118 

inconitus, 118 
Dasypodidae, 101 
Dasyprocta 

callida. 130 

colombiana, 130 

leporina, 130 

noblei, 130 

nuchalis, 130 

punctata, 130 
Dasyproctidae, 130 
Dasypus 

hoplites, 101 

novemcinctus, 101 
Dasyuiidae, 100 
Das)airomoi"phia, 100 
Daubentonia 

n ladagasca riens is , 135 
Daubentoniidae, 135 
davidanus 

Sciurotamias, 106 
decaryi 

Microgale, 102 
decUvis 

Beluga, 148 
degener 

Lutra, 152 
delectonim 

Praomys, 119 
deletrix 

Vidpes, 150 
Delphinapterus 

leucas, 148 
Delphinus 

nielas, 148 



demidovii 

Galago, 135 
Dendromurinae, 117 
desmarestianus 

Heteromys, 111 

Oryzomys, 123 
diadeina 

Propithecus, 135 
Dicrostonyx 

chionopaes, 113 

exsid, 113 

groenlandicus, 113 

torquatus, 113 
Didelphidae, 100 
Didelphis 

marsupialis, 100 

particeps, 100 

pigra, 100 

virginiana, 100 
Didelphimorphia, 99 
Diplomys 

labilis, 131 
Dipodidae, 112 
Dipodomys 

californicus. 111 

ordii. 111 

pallidulus. 111 

palmeri. 111 
Dipodops 

ordii. 111 

palmeri. 111 
dispar 

Sorex, 139 
distincta 

Neotoma, 122 
dorcfls 

Damaliscus, 149 
dorsalis 

Tamias, 109 
dorsatum 

Erethizon, 130 
Dremomys 

flavior, 104 

pemyi, 104 

se?iex, 104 
droidtardi 

Microgale, 102 
douglasii 

Tamiasciunis, 109 
dupreanum 

Eidolon, 98 
Echimyldae, 131 
Echyniipera 

australis, 101 

kalabu, 100 

rufescens, 101 
Eidolon 

dupreanum, 98 

sakalava, 98 
eZegans 



Spennophilus, 108 
elongata 

Mephitis, 164 

Proct/on, 151 
Emballonuridae, 142 
energuinenos 

Mustela, 153 

Putorius, 153 

Microtus, 115 
Eonycteris 

glandifera, 140 

spelaea, 140 
Eothenomys 

aurora, 113 

eua, 113 

melanogaster, 113 

mucronatus, 113 
Epimys 

zappeyi, 119 
Eptesicus 

darlingtoni, 145 

pliasma, 146 

pumilus, 145 
erem/cus 

Hesperomys, 98 

Peromyscus, 98, 126 
Erethizon 

dorsatum, 130 

picinus, 130 
Erethizontidae, 130 
erigens 

Hipposideros, 142 
Erioryzomys 

monochromos, 128 

Mustela, 152, 153 
Erophylla 

sezekomi, 144 

syops, 144 
erythraeus 

Callosciurus, 103 
Euarctos 

americanus, 151 

sornhorgeri, 151 
Eulipotyphla, 97 
Eu pie res 

goudotii, 155 

major, 155 
Eutamias 

minimus, 109 

neglectus, 109 
ewxmus 

Apod emus, 117 
eufl 

Eothenomys, 113 
Evotomys 

proteus, 113 
exsputus 

Sigmodon, 127 



174 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



exsul 

Dicrostonyx, 113 
extinius 

Sciurus, 106 
familiaris 

Cants, 150 
fasciatus 

Cervaria, 155 

Lynx, 155 
fatuus 

Synaptomys, 116 
Felidae, 155 
Fells 

bangsi, 156 

capensis, 155 

concolor, 156 

coryi, 156 

costaricensis, 156 

floridana, 156 

improcera, 156 

philUpsi, 155 

serval, 155 
femurvillosu m 

Artibeus, 144 
ferrugineus 

Callosciunis, 103 
feroidus 

Sigmodon, 128 
Fiber 

aquilonius, 116 

obscunis, 116 

rivalicius, 116 

zibethicus, 116 
fischeri 

Haplonycteris, 141 
flavicans 

Oecomys, 122 

Oryzomys, 122 
flavidus 

Isthniomys, 121 

Megadontomys, 121 
flavior 

Dremomys, 104 
flavipes 

Tat era, 117 
flaviventer 

Arctomys, 105 
flaviventris 

Marmot a, 105 
floridana 

Felis, 156 

Neotoma, 122 
floridanus 

Geomys, 110 

Sylvilagus, 134 
fontigenus 

Microtus, 115 
fontoynonti 

Dasogale, 103 
frenata 

Mustela, 152, 153, 154 



frenatus 

Putorius, 152 
fidvescens 

Oligoryzomys, 122 
fulviventer 

Mannosa, 99 
fumatus 

Myoniys, 119 

Praomys, 119 
Fimisciurus 

akka, 104 

pyrrhopiis, 104 

victoriae, 104 
furcuhis 

Triaenops, 143 
furvus 

Sigmodon, 128 

Urocyon, 150 
fuscipes 

Neotoma, 98 
fuscomurina 

Crocidura, 137 
fusus 

Peromyscus, 126 
Galago 

demidovii, 135 

orimis, 135 
Galagoides 

orinus, 135 
Galagonidae, 135 
Galidictis 

grandidiensis, 155 

grandidieri, 155 

Clethrionomys, 113 
geata 

Crocidura, 138 

Myosorex, 138 
Geocapromys 

abaconis, 132 

colutnbianus, 132 

cubanus, 132 

ingrahami, 132 

irrectus, 132 
GeogaZe 

aurita, 101 

orientalis, 101 
Geomyidae, 110 
Ggoinys 

austriniis, 110 

colonus, 110 

cumberlandius, 110 

floridanus, 110 

go#, 110 

pinetis, 110 

fwz.a, 110 
Gerbillinae, 117 
gerhillus 

Leggada, 119 
germanus 

Cerdocyon, 150 



gibbsi 

NeurotricJius, 140 
giganteiis 

Pteropus, 141 
gtgas 

Lynx, 155 
glaber 

Heterocephalus, 129 
glandifera 

Eonycteris, 140 
Glaucomys 

bangsi, 104 

lascivus, 104 

makkovikensis, 104 

querceti, 105 

sabrinus, 104 

uoZans, 104, 105 
Glossophaga 

longirostris, 144 
gloveralleni 

Procyon, 151 

Sorex, 138 
goffi 

Geomys, 110 
gorgonag 

Proechimys, 131 
gorilla 

Gorilla, 136 

Troglodytes, 136 
Gorilla 

gorilla, 136 
gossypinus 

Hesperomys, 123 

Peromyscus, 123, 124, 
125 
goudotii 

Eupleres, 155 
Grammonujs 

macmillani, 118 
granatensis 

Sciurus, 106, 107 
grandidiensis 

Galidictis, 155 
grandidieri 

Galidictis, 155 
grandis 

Orthogeonujs, 111 
Graphiurus 

argenteus, 129 

griseus, 129 

/lue^i 129 

lorraineus, 129 

microtis, 129 

murinus, 129 

schwabi, 129 

surdiis, 129 
griseus 

Graphiunis, 129 
griswoldi 

Tana, 134 
Guerlinmietus 



aestuans, 106 

chiriquensis, 106 
guyannensis 

Proechimys, 131 
gymnicus 

Sciurus, 109 

Tamiasciunis, 109 
haemobaphes 

Callosciunis, 103 

Sciurus, 103 
Haplonycteris 

fischeri, 141 
hardyi 

Zapus, 112 
haiyia 

Harpiocephalus, 146 
Hai~piocephalus 

harpia, 146 

nifuhis, 146 
harringtoni 

TateriUus, 117 
helleri 

Platyrrhinus, 144 
Herpestidae, 155 
hesperia 

Lonchophylla, 144 
Hesperomys 

arcticus, 124 

eremicus, 98 

gossypinus, 123 

leucopus, 124 

nebrascensis, 124 

sonoriensis, 124 
Heterocephalus 

glaber 129 

stygius, 129 
Heteromyidae, 111 
Heteromys 

alleni, 112 

desmarestianus. 111 

repens. 111 
hildegardeae 

Crocidura, 138 
Hipposideros 

bicolor, 142 

curtus, 142 

erigens, 142 

turpis, 143 
hirsutiis 

Sigmodon, 128 
hispid a 

Suillomeles, 100 
hispidus 

Sigmodon, 127, 128 
Hominidae, 138 
hoplites 

Dasypus, 101 
hottentotus 

Cryptomys, 129 
hudsonica 

Lutra, 152 



Type Specimens of Recent Mammals • Helpen and McFadden 175 



luidsonlciis 

Sciunis, 109 

Tainiasciurus, 109 

Zapus, 112 
htieti 

Aedioglis, 129 

Graphiurus, 129 
huinidis 

Reifli rodoiifoinys, 127 
hyacindiits 

Neiirotriclius, 140 
hijleae 

Proechimijs, 131 
Hylobates 

carpenteri, 136 

lai; 136 
Hylobatidae, 136 
Hijlomyscus 

alleni, 118 

simus, 118 
Hijlopetes 

alboniger, 105 

anchises, 105 

oriniis, 105 

phaijrei, 105 
hypoxanthus 

Oenomys, 119 
Hypudaeus 

nivicola, 112 
Ictidomys 

badius, 108 

trideceinllneatus, 108 
Idionycteris 

phyllotis, 146 
ignaua 

Marmot a, 105 
jgnfluxs 

Arctomys, 105 
igni/er 

Rhinolophus, 143 
illectiis 

Oecotm/s, 122 

Oitjzomys, 122 
impiger 

Reldirodontoinys, 127 
improcera 

Fells, 156 

Puma, 156 
incitatus 

Lepus, 134 

Sylvilagus, 134 

Tapeti, 134 

Dasymys, 118 
Indridae, 135 
ingrahami 

Geocapromys, 132 
innultus 

Mictomys, 116 

Synaptomys, 116 
insulanus 



Peromyscus, 125 
insularis 

Pteropus, 98 

Geocapromys, 132 
irroratus 

Liomys, 112 
Isdimomys 

flavidus, 121 
kalabit 

Echymipera, 100 

T«f<?ra, 117 
kivtiensis 

Thamnomys, 120 
kodiacensis 

Spennopliilus, 99 

Microtus, 115 

Sorex, 138 
labilis 

Diplomys, 131 

Loncheres, 131 
lacustris 

Otomxjs, 120 
ladas 

Zapus, 112 
laetus 

Aethosciunis, 106 
Lagomoi'plia, 133 
lambertoni 

Nesomys, 120 
lanatus 

Cahiromys, 99 
laniger 

Cheiromys, 135 

Pteropus, 98 
lanigera 

Pteropus, 98 

Hylobates, 136 
lascivus 

Glaucomys, 104 

Sciuroptenis, 104 
Lasiurus 

borealis, 146 

brachyotis, 146 
latimanus 

Scapanus, 140 
laxatina 

Lontra, 142 
lecontii 

Reithrodontomys, 127 
Leggada 

ablutus, 118 

fou/o, 118 

gerhtllus, 119 
Lemr7ius 

chrysogaster, 114 

paulus, 114 

sibiricus, 114 



lepida 

Neotoma, 121 
Leporidae, 143 
leporina 

Dasyprocta, 130 
Leptailunis 

phillipsi, 155 

serval, 155 
LepM5 

alacer, 134 

alleni, 133 

americanus, 133, 134 

arcticus, 133 

Z?air<ii, 133, 134 

bangsii, 133 

cascadensis, 134 

incitatus, 134 

palitans, 133 

paludicola, 134 

stnithopus, 133 

sylvaticus, 134 

transltionalis, 134 

Delphinaptenis, 148 
leucogaster 

Scotophilus, 147 
leucopus 

Hesperomys, 124 

Peromyscus, 126 
leucostigma 

Chaerophon, 145 

Mops, 145 
levipes 

Melomys, 118 
hiomys 

alleni, 112 

irroratus, 112 
Lipotyphla, 136 
lituratus 

Artibeus, 144 
Loncheres 

labilis, 131 
Lonchophylla 

hesperia, 144 
longicauda 

Arctogale, 153 

Putorius, 153 
longicaudata 

Microgale, 102 
longimembis 

Perognathus, 112 
longipilis 

Arvicola, 114 
longirostris 

Glossophaga, 144 
Lonfra 

canadensis, 152 

laxatina, 152 

Sciurus, 109 
Tamiasciurus, 109 



lorraineus 

Graphiurus, 129 
Zotor 

Procyon, 151 
louisianae 

Odocoileus, 148 
lucifugus 

My Otis, 146 
luctuosa 

Alouatta, 135 
ludovicianus 

Sciurus, 106 
lupus 

Canis, 149, 150 
lutensis 

Lutreola, 153 

Mustela, 153 

Putorius, 153 

degener, 151 

hudsonica, 151 

uago, 151 
Lutreola 

lutensis, 153 

vulgivagus, 154 
Lynx 

calif ornicus, 155 

canadensis, 155, 156 

fasciatus, 155 

gigas, 155 

oculeus, 155 

nifus, 155 

subsolanus, 156 
mabirae 

Phataglnus, 156 
macmillani 

Grammomys, 118 
Macrogeomys 

cavator, 110 

pansa, 110 
macropygmaeus 

Sorex, 138 
macrotis 

Nycteris, 142 

Peropteryx, 142 
Macrotolagus 

alleni, 133 

palitans, 133 
macrou ra 

Odocoileus, 148 
macnirus 

Sorex, 139 
madagascariensis 

Cheiromys, 135 

Daubentonia, 135 

Nycteris, 142 

Rousettus, 141 
major 

Eupleres, 155 

Vampyrodes, 145 
makkovikensis 



176 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Glaucomys, 104 

Sciuroptenis, 104 
maniculatus 

Peromyscus, 98, 124, 
125 
Manidae, 156 
marcanoi 

Antillogale, 137 

Solenodon, 137 
Marmosa 

fulviventer, 99 

mitts, 99 

robinsoni, 99, 100 
Marmosidae, 99 
Marmota 

avara, 105 

flaviventris, 105 

ignava, 105 

monax, 105 
inarsupialis 

Didelphis, 100 
Martes 

americana, 152 

atrata, 152 

bnimalis, 152 
maurisca 

Crocidura, 138 
mayeri 

Antechinus, 100 
rnaynardi 

Procyon, 151 
meamsz 

Saccostomus, 117 
niedialis 

Callosciunis, 103 
niedioximii.s 

Mictomys, 116 

Synaptomys, 116 
Megadontomys 

flavidus, 121 
megaphyllus 

Rhinolophus, 143 
melanogaster 

Eothenomys, 113 
melanops 

Taterillus, 117 
melanotis 

Oryzomys, 123 
melanotus 

Praomys, 119 
meZas 

Delphinus, 148 

Myoictis, 100 
Melornys 

alleni, 118 

levipes, 118 

mollis, 118 

moncktoni, 118 

ruhex, 118 

stevensi, 118 
mephitica 



Mephitis, 154 
Mephitidae, 154 
mephitis 

Mephitis, 154 
Mephitis 

avia, 154 

elongata, 154 

mephitica, 154 

mephitis, 154 

mesomelas, 154 

scrutator, 154 

spissigrada, 154 
Mesocap romys 

nanus, 132 
mesomelas 

Mephitis, 154 

Neotoma, 122 
mexicanus 

Peromyscus, 124 
Microgale 

brevicaudata, 102 

cowani, 102 

decanji, 102 

droiihardi, 102 

longicaiidata, 102 

parvula, 102 

principtda, 102 

prolixicaudata, 102 

pu/Zfl, 102 
Microsciurus 

alfan, 105 

browni, 105 
microtis 

Graphiuriis, 129 
Microtus 

acadtcus, 115 

aurora, 113 

breweri, 114 

chrotorrhinus, 1 14 

enixus, 115 

fontigenus, 115 

koreni, 115 

mucronatus, 113 

oeconomus, 115 

pennsylvanicus, 114, 
115 

provectus, 115 

ravus, 114 

shattucki, 115 

terraenovae, 114 

Nesophontes, 136 
Mictomys 

innuitus, 116 

medioximus, 116 
minimus 

Tamias, 99, 109 

Ametrida, 143 
Suncus, 139 



minusculus 

Scapanus, 140 
mjsczx 

Sorex, 139 
misim 

Antechinus, 100 
mitis 

Marmosa, 99 
mollipilosus 

Tamiasciunis, 109 
mollis 

Melonujs, 118 

Stenomys, 120 
Molossidae, 145 
monax 

Marmota, 105 
moncktoni 

Melomys, 118 
monochromos 

Erioryzomys, 128 

Oryzomys, 128 

Thomasomys, 128 
Monodontidae, 148 
monoensis 

Pteropus, 141 
Mops 

angolensis, 145 

condylunis, 145 

leucostigma, 145 

orientis, 145 
Mormoopidae, 143 
mortigena 

Mustela, 152 
morulus 

Sciurus, 107 
mucronatus 

Microtus, 113 
munda 

Mustela, 154 
mundus 

Putorius, 154 
muricola 

Myotis, 146 
initnctis 

Arctogale, 153 

Mustela, 153 

Putorius, 153 
Muridae, 112 
Murinae, 117 
murinus 

Graphiuriis, 129 

Pseudohydromys, 120 
Mms 

bactrianus, 119 

bairdii, 98 

Z;r//o, 118 

niusculus, 119 

tantillus, 119 

tenellus, 119 
musculiis 

Mus, 119 



Mustela 

atrata, 152 

bnimalis, 152 

cicognanii, 152 

energiimenos, 153 

erminea, 152, 153 

frenata, 152, 153, 154 

lutensis, 153 

mortigena, 152 

munda, 154 

muricus, 153 

neomexicana, 152 

nivalis, 153 

noveboracensis, 153 

occisor, 153 

oribasus, 153 

richardsonii, 152 

rixosa, 153 

vison, 153, 154 

vulgivaga, 154 
Mustelidae, 152 
Myoictis 

nielas, 100 

wallacei, 100 

wavicus, 100 
Myomys 

funmtus, 119 
Myopus 

schisticolor, 115 

thayeri, 115 
Mi/osorex 

geata, 138 
Myofjs 

abbotti, 146 

albicinctus, 146 

carissima, 146 

lucifugus, 146 

muricola, 146 

nugax, 146 

sodalis, 146 
Myoxidae, 129 
mystacinus 

Apodemus, 117 
nana 

Capromys, 132 

Nycteris, 142 
nanus 

Mesocapromys, 132 
nrtso 

Antechinus, 100 

Rhynchiscus, 142 
natator 

Oryzomi/s. 123 
naiyws 

Oryzomi/s, 122 
nebrascensts 

Hesperomys, 124 

Peromyscus, 124 
Nectomys 

aniazonicus, 121 

squamipes, 121 



Type Specimens of Recent Mammals • Helgen and McFadden 111 



neglectus 

Tainias, 109 
Neomeris 

asiaeorientalis, 148 
neomexicana 

Musfelfi, 152 
neoinexica)}its 

Putoriiis, 152 
Neophocaena 

asiaorientalis, 148 

phocaenoides, 148 
Neosoi-ex 

acadicus, 138 

palttstris, 138 
Neotonia 

abhreviata, 121 

beUa, 121 

distincta, 122 
floridana, 122 

fuscipes, 98 

lepida, 121 

mexicana, 122 

riibida, 122 
nesaeiis 

Scitinis, 107 
Nesomylnae, 120 
Nesomys 

lambertoni, 120 
Nesophontes 

micriis, 136 
Nesophontidae, 136 
Neil rot richits 

gibbsi, 140 

Jiyaciiidihiiis, 140 

Sciiiriis, 106 
nigrictihis 

Peromysciis, 125 

StenoDiys, 120 
nitellinits 

Nyctoinys, 122 
nivalis 

Chionomys, 112 

Mustela, 153 
nivicola 

Hypudaeiis, 112 
Niviventer 

confuciainis, 119 
noblei 

Dasyprocta, 130 

Pipistrelliis, 147 

Putorius, 153 

Scabrifer, 147 
noveboracensis 

Mustela, 153 

Putorius, 153 
novemcinctus 

Dasypus, 101 
nuchalis 



Dasyprocta, 130 
nugax 

My Otis, 146 

Steatomys, 117 
Nycteridae, 142 
Nycteris 

macrotis, 142 

madagascariensis, 142 

nana, 142 

revoili, 97 

tristis, 142 
Nycticeiiis 

africanus, 147 

albiventer, 147 

schlieffeni, 147 
Nyctomys 

nitellinus, 122 

sumichrasti, 122 
ohsciinis 

Citellus, 107 

FAer, 115 

Ondatra, 115 
obsoletus 

Spennophilus, 99 
occidentalis 

Paramicrogale, 102 
occisor 

Mustela, 153 

Putorius, 153 
occlusus 

Cryptonu/s, 129 
Ochotona 

cupppes, 133 

princeps, 133 

saxatilis, 133 
Ochotonidae, 133 
ochraceiis 

Thamnomys, 118 
Octodontidae, 130 
Octomys 

barrerae, 130 

Cervaria, 155 

Lynx, 155 
ocythoiis 

Urocyon, 150 
Odocoileus 

americanus, 148 

borealis, 148 

clavium, 149 

louisianae, 146 

macroura, 146 

osceola, 146 

virginianus, 146, 147 

bicolor, 122 
flavicans, 122 
illectus, 122 
trabeatus, 122 
oecono»?i;<s 



Microtus, 115 

Oenomys 

hypoxanthus, 119 
talangae, 119 

Oligoryzornys 
fulvescens, 122 
vegetus, 123 

Ondatra 

ac/uilonius, 116 
obscunis, 116 
rivalicius, 116 
zibethictis, 116 

orariiis 

Sciunis, 109 
Zapus, 112 

ordii 

Dipodomys, 111 
Dipodops, 111 

oreas 

Peromyscus, 126 

oribasus 

Arctogale, 153 
Mustela, 153 
Putorius, 153 

orientalis 

Geogale, 101 

orientis 
Mops, 145 

onn«.S' 

Gala go, 135 
Galagoides, 135 
Hylopetes, 105 
Pteromys, 105 

Orthogeoniys 
cavator, 110 
grandis. 111 
pansa, 110 
p/((fo. 111 

Oryzoinys 

carrorum, 123 
coloratus, 123 
devius, 123 
flavicans, 122 
fulvescens, 122 
illectus, 122 
nielanotis, 123 
monochronios, 128 
natator, 123 
navus, 122 
palustris, 123 
rostratus, 123 
vegetus, 123 

osceoZa 

Cariacus, 148 
Odocoileus, 146 

Otomops 

papuensis, 145 

Otomyinae, 120 

anchietae, 120 
barboiiri, 121 



lacustris, 120 
typus, 121 
uzungwensis, 121 

Praoinys, 119 

Agouti, 130 
Pachycyon 

robustus, 150 
paitana 

Tupaia, 134 
palitans 

Lepus, 133 

Macrotolagus, 133 
palliata 

Alouatta, 135 
pallidulus 

Dipodomys, 111 
pallidus 

Spennophilus, 99 

Tainias, 99 
pahnarius 

Peromyscus, 125 
pabnarum 

Artibeus, 144 
pahneri 

Dipodomys, 111 

Dipodops, 111 
paludicola 

Lepus, 134 

St/lvilagus, 134 
palustris 

Neosorex, 138 

Onjzomys, 123 

Sorex, 138 

Sylvilagus, 134 
pansa 

Macrogeomys, 1 10 

Ortliogeonujs, 110 
Pappogeomys 

castanops. 111 

ntbellus. 111 
papuensis 

Otomops, 145 
Paracaproniys, 132 
Paramicrogale 

occidentalis, 102 
Paraxenis 

byatti, 106 

vexillarius, 106 
paniellii 

Chilonycteris, 143 

Pteronotus, 143 
parryi 

Spennophilus, 99 
particeps 

Didelphis, 100 
parvula 

Microgale, 102 
paulus 

Leninuis, 114 



178 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2 



Pecari 

ci-usnigrum, 147 

tajacu, 147, 148 

torvus, 148 
Pedetes 

cafer, 128 

capensis, 128 

taborae, 128 
Pedetidae, 128 
pennsylvanica 

Vulpes, 150, 151 
pennsylvanicus 

Microtus, 114, 115 
Peramelia, 100 
pernyi 

Dremomys, 104 
Perognathus 

hangsi, 112 

longiinembris, 112 
Peromyscus 

abietoruin, 125 

ammodytes, 125 

anastasae, 124, 125 

arenarius, 126 

argentatus, 125 

attwateri, 124 

bairdii, 98 

baliolus, 126 

foeZZtis, 124 

borealis, 124 

cacabatiis, 124 

canadensis, 125 

crinitus, 125 

eremicus, 98, 126 

fusus, 126 

gossypinus, 123, 124, 
125 

insidanus, 125 

leucopus, 126 

maniculatus, 98, 124, 
125 

mexicaniis, 124 

nebrascensis, 124 

nigriculus, 125 

oreas, 126 

pahnarius, 125 

phasma, 126 

polionotus, 126 

rhoadsi, 126 

saturatus, 126 

scitulus, 125 

subgrisetis, 126 

texanus, 126 
Peropteryx 

canina, 142 

macrotis, 142 

phaea, 142 
Peroiyctidae, 100 
perneri 

Propithecus, 135 
personatus 



Sorex, 139 
phaea 

Peropteryx, 142 
phaios 

Crocidura, 138 
phasma 

Eptesicus, 147 

Peromyscus, 126 

Pipistrellus, 147 
Phataginus 

niabirae, 156 

tricuspis, 156 
phayrei 

Hylopetes, 105 

Pterornys, 105 
Phenacomys 

celatus, 116 

crassus, 116 

ungava, 116 
Philander 

ciciir, 99 
philippinensis 

Rhinolophus, 143 
phillipsi 

Damaliscus, 149 

Fe/is, 155 

Leptatlunis, 155 
phocaenoides 

Neophocaena, 148 
Phocoenidae, 148 
Phyllostomidae, 143 
phyllotis 

Corynorhinus, 146 

Idionycteris, 146 
picnii/s 

Erethizon, 130 
pigra 

Alouatta, 135 

Didelphis, 100 
piloroides 

Capromys, 131 
pinetis 

Geomys, 110 
Pipistrellus 

capensis, 147 

notius, 147 

phasma, 147 

rendalli, 147 
Plagiodonta 

araeum, 132 
Platyrrhinus 

helleri, 144 

umbratus, 144 
Plecotus 

aiiritus, 147 

sacrimontis, 147 
pluto 

Orthogeomys, 111 
poeyanus 

Solenodon, 137 
polionotus 



Peromyscus, 126 
Praomys 

delectorum, 119 
fumatus, 119 

melanotus, 119 

oweni, 119 

tuHbergi, 119 
pratensis 

Steatomys, 117 
Primates, 135 
primus 

Callosciimts, 103 
pnuceps 

Ochotona, 133 
principula 

Micro gale, 102 
prisons 

Rhynchiscus, 142 
Procyon 

elucus, 151 

gloveralleni, 151 

Zofor, 151 

maynardi, 151 
Procyonidae, 151 
Proechimys 

burnis, 131 

cayennensis, 131 

gorgonae, 131 

guyannensis, 131 

hyleae, 131 

semispinosus, 131 
p rolixicaudata 

Microgale, 102 
Propithecus 

diadema, 135 

perrieri, 135 
proteus 

Clethrionomys, 113 

Evotomys, 113 
provectus 

Microtus, 115 
Pseudohydromys 

murinus, 120 
Pferojnys 

alboniger, 105 

anchises, 105 

orinus, 105 

phayrei, 105 
Pteronotus 

pamellii, 143 

pusillus, 143 

quadridens, 143 
Pteropodidae, 140 
Pteropus 

anetianus, 141 

aorensis, 141 

rtrie/, 141 

austini, 141 

giganteus, 141 

insidaris, 98 

laniger, 98 



lanigera, 98 

monoensis, 141 

rayneri, 141 

woodfordi, 141 
pulla 

Microgale, 102 
Pu??ifl 

hangsi, 156 

concolor, 156 

coryi, 156 

costaricensis, 156 

improcera, 156 
punctata 

Dasyprocta, 130 
pusillus 

Chilonycteris, 143 

Pteronotus, 143 
putorius 

Spilogale, 155 
Putorius 

energumenos, 153 
frenatus, 152 

longicauda, 153 

lutensis, 153 

mundus, 154 

muricus, 153 

neomexicanus, 153 

notius, 153 

noveboracensis, 153 

occisor, 153 

oribasus, 153 

rixosus, 153 

vison, 153 

vidgivagus, 154 

xanthogenys, 154 
pj/gargMS 

Damaliscus, 149 
pyrrhopus 

Funisciunis, 104 
quadridens 

Pteronotus, 143 
qiiadrivittatus 

Tamias, 99, 109 
querceti 

Glaucomys, 105 

Sciuropterus, 105 
ramieri 

Aplodontia, 103 
Rangifer 

arcticus, 149 

caboti, 149 

caribou, 149 

tarandus, 149 

terraenovae, 149 
rat>u.s 

Microtus, 114 
rayneri 

Pteropus, 141 
Reithrodontomys 

australis, 127 

creper, 127 



Type Specimens of Recent Mammals • Helaen and McFadden 179 



hinutills, 127 

itnpiger, 127 

lecontii, 127 

sumichrasti, 127 

viilcaniiis, 127 
relicttis 

Capromys, 131 
rendalli 

Pipistrelhis, 147 
repens 

Heteroimjs, 111 

Nijcteris, 97 
Rhinolophidae, 142 
Rliinolopliiis 

alleni, 143 

ig;nifer, 143 

megaphyllus, 143 

philippinensis, 143 
rhoadsi 

Peroinijscus, 126 
Rhynchiscus 

naso, 142 

priscus, 142 
RJn/nchonycteris 

naso, 142 
richardsonii 

Mustela, 152 
riparia 

Arvicola, 114 
rivalicius 

Fiber, 116 

Ondatra, 116 
rixosrt 

Mustela, 153 
nxosrts 

Piitorius, 153 
robinsoni 

Mannosa, 99, 100 
roboratus 

Sorex, 139 
robustiis 

PacJu/cyon, 150 
Rodentia," 103 
ro stratus 

Oryzomys, 123 
Rousettus 

madagasca riens is , 141 
nibellus 

Cratogeomys, 111 

Pappogeoniys, 111 
nibex 

Melomys, 118 
rubida 

Neotoma, 122 
rubricatus 

Vulpes, 151 
nibricosa 

Vulpes, 150, 151 

Aplodontia, 103 



nifescens 

Echymipera, 101 
rufidorsunx 

Arvicola, 114 
rufulus 

Haiyioceplialus, 145 
rufus 

Lynx. 155 
sabrinus 

Glaucoinys, 104 

Sciuroptenis, 104 
Saccostomus 

cricetulus, 117 

meamsi, 117 
sacrimontis 

Plecotus, 147 
sakalava 

Eidolon, 98 
saltuensis 

Sciunis, 107 
sanctaeniartae 

Sigmodon, 128 
sanguinidens 

Sorex, 139 
saturatus 

Peroniyscus, 126 
saxatilis 

Ochotona, 133 
Scabrifer 

notius, 147 
Scalops 

aereus, 140 

anastasae, 140 

texanus, 140 
Scalopus 

aereus, 140 

anastasae, 140 

aquaticus, 140 
Scandentia, 134 
Scapanus 

califomicus, 140 

latirnanus, 140 

minusculus, 140 
schisticolor 

Myopus, 115 
schliejfeni 

Nycticeius, 147 
Schwab i 

Graphiurus, 129 
scitulus 

Peroniyscus, 125 
Sciuridae, 103 
Sciuroptenis 

alpinus, 104 

bangsi, 104 

lascivus, 104 

niakkovikensis, 104 

querceti, 105 

sabrinus, 104 

.siZiis, 104 

ooZons, 104, 105 



Sciurotaniias 

consobrinus, 106 

davidianus, 106 

thayeri, 106 
Scit<n<s 

aberti, 99 

aestuans, 106 

browni, 105 

carolinensis, 106 

castaneoventris, 103 

castanonotus, 99 

castanotus, 99 

chiriquensis, 106 

extimus, 106 

granatensis, 106, 107 

gymnicus, 109 

haemobaphes, 103 

hudsonicus, 109 

loquax, 109 

ludovicianus, 106 

mondus, 107 

nesaeus, 107 

niger, 106 

orarius, 109 

saltuensis, 107 

variabilis, 107 

vicinus, 106 

vulpinus, 106 

ijucatanensis, 106 
Scotinomys 

apricus, 127 

teguina, 127 

xeranipelinus, 127 
Scotophilus 

altilis, 147 

leucogaster, 147 
scrutator 

Mephitis, 154 
seniispinosus 

Proechimys, 131 
senex 

Drenionujs, 104 

Zygodontomys, 128 
serval 

Felis, 155 

Leptailunis, 155 
Setifer 

setosus, 103 
setosus 

Setifer, 103 

Erophylla, 144 
shattucki 

Microtus, 115 
sibiricus 

Lemrnus, 114 
siccus 

Citellus, 107 
Sigmodon 

austendus, 127 



borucae, 127 
exsputus, 127 
fervidus, 128 
furvus, 128 
hirsutus, 128 
hispidus, 127, 128 
sanctaeniartae, 128 
spadicipygus, 128 

Sigmodoiitinae, 121 

siZus 

Sciuroptenis, 104 

SiHUiS 

Hylomyscus, 118 
sodalis 

Myotis, 146 
soleatus 

Claviglis, 129 
Solenodon 

cubanus, 137 

marcanoi, 137 

poeyanus, 137 
Solenodontidae, 137 
sonoriensis 

Hesperomys, 124 
Sorex 

araneus, 138 

caecutiens, 138 

cinereus, 139 

daphaenodon, 139 

dispar, 139 

gloveralleni, 138 

koreni, 138 

inacropygniaeus, 138 

niaci~urus, 139 

miscix, 139 

palustris, 138 

personatus, 139 

roboratus, 139 

sanguinidens, 139 

tundrensis, 138 

ultinuis, 138 

oir, 139 
Soricidae, 137 
sornborgeri 

Euarctos, 151 

L/rs((s, 151 
soro/- 

Trtfera, 117 
spadicipygus 

Sigmodon, 128 
spelaea 

Eonycteris, 140 
Spennophilus 

alashanicus, 107 

arniatus, 107 

badius, 108 

elegans, 108 

kodiacensis, 99 

obsoletus, 99 

pallidus, 99 

parryi, 99 



180 Bulletin Museum of Comparative Zoology, Vol. 157, No. 



spilosoma, 99 

texensis, 108 

tridecemlineatus, 99, 
108 
Spilogale 

ainbarvalis, 155 

putorius, 155 
spissigrada 

Mephitis, 154 
squarnipes 

Nectomys, 121 
Steatomys 

nyasae, 117 

pratensis, 117 
Stenoinys 

mollis, 120 

niobe, 120 

stevensi, 120 

verecundus, 120 

Melomys, 118 

Stenoinijs, 120 
striatus 

Tamias, 107 
stnithopus 

Lepiis, 133 
stuhhnanni 

Chnjsochloris, 101 
stygius 

Heterocephalus, 129 
subgriseus 

Peromyscus, 126 
suhsolanus 

Lynx, 156 
Suillomeles 

hispida, 100 
sumichrasti 

Nyctomys, 122 

Reithrodontomys, 127 
Suncus 

ater, 139 

minor, 139 

varilla, 139 

Graphiunis, 129 
sylvaticus 

Lepus, 134 
Sylvilagus 

alacer, 134 

brasiliensis, 134 
floridanus, 134 

incitatus, 134 

paludicola, 134 

palustris, 134 

transitionalis , 134 
Synaptomys 

boreal is, 116 

cooperi, 116 
fatuus, 116 

innuitus, 116 

medioxitnus, 116 



Syntheosciwins 

broclius, 108 
syops 

Erophylla, 144 
taborae 

Pedetes, 128 
tajacu 

Pecari, 147, 148 
talangae 

Oenomys, 119 
Talpidae, 140 
Tamias 

borealis, 109 

cooperi, 108 

dorsalis, 108 

minim.us, 99, 109 

neglectus, 109 

pallidus, 99 

quadrivittatus, 99 

striatus, 109 

townsendii, 108 

venustus, 109 
Tamiasciunis 

douglasii, 109 

gym,nicus, 109 

hudsonicus, 109 

loquax, 109 

mollipilosus, 109 
Tanfl 

grisiooldi, 134 

tona, 134 
fana 

Tana, 134 

Tupaia, 134 
tantilliis 

Mas, 119 

incitatus, 134 
ta rand us 

Rangife7', 149 
Tafera 

flavipes, 117 

kempi, 117 

soror, 117 

valida, 117 
Taterillus 

liarringtoni. 117 

melanops, 117 
Tayassu 

crusnignim, 147 

torvus, 148 
Tayassuidae, 147 
teguina 

Akodon, 127 

Scotinomys, 127 
tenellus 

Mus, 119 
Tenrecidae, 101 
tephragaster 

Crocidura, 137 
terraenovae 



A)~vicola, 114 

Microtus, 114 

Rangifer, 149 
texanus 

Peromyscus, 126 

Scalops, 140 
fexe'n.sz.s 

Spernwpliilus, 108 
Thamnomys 

kivuensis, 120 

ochraceus, 118 

venustus, 120 
thayeri 

My opus, 115 

Sciurotamias, 106 
thoniasi 

Crypt Otis, 138 
Thomasomys 

nionochromos, 128 
Thoniomys 

atrodorsalis. 111 

unibrinus. 111 

Cerdocyon, 150 
Thyroptera 

albigula, 147 

alhiventer, 147 

tricolor, 147 
Thyropteridae, 147 
torquatus 

Dicrostonyx, 113 
torrei 

Boromys, 131 

Chilonycteris, 143 
toruw5 

Pecari, 148 

Tayassu, 148 
townsendii 

Tamias, 109 
trabeatiis 

Alouatta, 135 

Oecomys, 122 
transitionalis 

Lepus, 134 

Sylvilagus, 134 
Triaenops 

aurita, 143 
furculus, 143 
fricoZor 

Thyroptera, 147 
tricuspis 

Phataginus, 156 
tridecemlineatus 

Spennophilus, 99, 108 
trinotatus 

Zapus, 112 

Nycteris, 142 
Troglodi/tes 

gorilla, 136 
tropicalis 



Chlorotalpa, 101 

Chnjsochloris, 101 
tullbergi 

Praomys, 119 
tundrensis 

Sorex, 138 
Tupaia 

paitana, 134 

tona, 134 
Tupaiidae, 134 
turpis 

Hipposideros, 143 

Geo my s, 110 
Tympanoctomys 

barrerae, 130 
typus 

Otomys, 121 
ultimus 

Sorex, 138 
umbratus 

Platyrrhinus, 144 

Vampyrops, 144 
umbrinus 

Thomonu/s, 111 
ungava 

Phenacomys, 116 
Urocyon 

aquilus, 150 

cinereoargenteus, 150 
furvus, 150 

ocythous, 150 
Ursidae, 151 

americanus, 151 

somborgeri, 151 
uzungwensis 

Otomys, 121 
vafra 

Vulpes, 150, 151 
uaga 

L«fra, 152 
valida 

Tatera, 117 
Vampijrodes 

caraccioli, 145 

major, 145 
Vampyrops 

umbratus, 144 

zarhinus, 144 
variabilis 

Sciurus, 107 
varilla 

Suncus, 139 
uege"f!/.s 

Oligoryzonujs, 123 

Oryzomys, 123 
venustus 

Tamias, 109 

Thamnomys, 120 
verecundus 



Type Specimens of Recent Mammals • Helpen and McFadden 



181 



Stenoinys, 120 
Vespertilionidae, 145 
vexillariiis 

Paraxerus, 106 
vicinus 

Sciunis, 106 
victoriae 

Fitnisciiinis, 104 
vir 

Sorex, 139 
virgatus 

Agouti, 130 
virginiana 

Didelphis, 100 
virginianus 

Odocoileus, 148, 149 
vison 

Mnstela, 153, 154 

Putoriiis, 153 



Viverridae, 155 
volans 

Glaucomys, 104, 105 

Sciuroptenis, 104, 105 
vidcanitis 

Reithrodoutomi/s, 127 
vidgivaga 

Mustela, 154 
vtdgivagus 

Lutreola, 154 

Putorius, 154 
vidpes 

Vtdpes, 150, 151 
Vidpes 

bnngsi, 151 

deletrix, 150 

pennsylvanica, 150, 151 

ruhricatus, 151 

nibricosa, 150, 151 



ufl/ra, 150, 151 

vulpes, 150, 151 
vulpinus 

Sciun.is, 106 
ivallacei 

Myoictis, 100 
wavicus 

Myoictis, 100 
whijtei 

Cryptomys, 129 
woodfordi 

Pteropus, 141 
xanthogenys 

Putorius, 154 
xerampelinus 

Akodon, 127 

Scotinoniys, 127 
yucatanensis 

Sciurus, 106 



zappeyi 

Epirnys, 119 

Zapus 

acadicus, 112 
luirdyi, 112 
hudsonicus, 112 
ladas, 112 
orarius, 112 
trinotatus, 112 

zarhinus 

Vampyrops, 144 

zibethicus 
Fiber, 116 
Ondatra, 116 

Zygodontomys 
brevicauda, 128 
cheniei, 128 
seorsus, 128 



(US ISSN 0027-4100) 



OF THE 



seum 



A New Species of Lizard Related to 
Stenocercus caducus (Cope) (Squamata: 
Iguanidae) From Peru and Bolivia, With a 

Key to the "Ophryoessoides Group" 



JOHN E. CADLE 


IJBRARY 




OCT 2 3 2002 




HARVARD 


HARVARD UNIVERSITY 

CAMBRIDGE, MASSACHUSEI IS, U.S.A. 


VOLUME 157, NUMBERS 
14 November 2001 



(US ISSN 0027-4100) 



PUBLICATIONS ISSUED 

OR DISTRIBUTED BY THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

HARVARD UNIVERSITY 



Breviora 1952- 

bulletin 1863- 

Memoirs 1865-1938 

JoHNSONiA, Department of Mollusks, 1941-1974 

Occasional Papers on Mollusks, 1945- 

SPECIAL PUBLICATIONS. 

1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and 
Evolution of Crustacea. 192 pp. 

2. Turner, R. D., 1966. A 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 pennanent paper stock. 

©The President and Fellows of Harvard College 2001. 



A NEW SPECIES OF LIZARD RELATED TO STENOCERCUS 
CADUCUS (COPE) (SQUAMATA: IGUANIDAE) FROM PERU AND 
BOLIVIA, WITH A KEY TO THE "OPHRYOESSOIDES GROUP " 



JOHN E. CADLE1 

CONTENTS 



Abstract -- 183 

Resumen 183 

Introduction 184 

Materials and Methods 184 

Description of a New Species of Stenocersus 187 

Stenocercus prionotus new species _ 187 

Diagnosis and Comparisons 191 

Description „ 193 

Distribution Patterns in Stenocercus prionotus 

197 

Patterns of Sympatry and Geographic Variation 

in Stenocercus prionotus, and the Need for 

Additional Fieldwork 200 

Natural Histoiy of Stenocercus prionotus 205 

Comparison of Stenocercus prionotus with S. 

caducus 206 

Distributions of Stenocercus prionotus and S. 

caducus in Eastern Bolivia 210 

Is the Distribution of Stenocercus aculeatus 

disjunct? 212 

Key to Species of the " Ophryoessoides group" 

of Stenocercus 212 

Acknowledgments 215 

Appendix: Specimens Examined 216 

Literature Cited 218 

Abstract. A new species of iguanid lizard, Steno- 
cercus prionotus, is described from eastern Peiii and 
Bolivia (known range froin San Martin Department, 
Peru, to northern La Paz and El Beni departments, 
Bolivia). Most localities are in the Andean foothills 
and immediately adjacent lowlands. Stenocercus 
prionotus is similar to several other species of Sten- 
ocercus with large posterior head scales, an enlarged 
row of supraoculars, and keeled ventral scales. These 
similar species are referred to as the "Ophryoessoides 
group" without implying that it is a monophyletic as- 
semblage. Based on their common possession of a 



' Department of Herpetology, Chicago Zoological 
Society, 3300 Golf Road, Brookfield, Illinois 60513. 
Research Associate, Department of Herpetology, 
Museum of Comparative Zoology. 



unique scaly flap concealing a portion of the posthu- 
meral mite pocket, the new species is apparently 
closely related to S. caducus (Cope), which is known 
from central Bolivia south to northern Argentina and 
Paraguay. Stenocercus prionotus is distinguished from 
S. caducus by having a more prominent vertebral 
crest and a pattern of alternating light and dark bars 
on the throat (rarely obseived in S. caducus, which 
usually has light throat spots). These two species also 
occupy different physiographic regions (western Am- 
azonian rainforest for S. prionotus; chaco for S. cad- 
ucus). 

Populations of S. prionotus from northern Peru 
have a higher vertebral crest tlian those in southern 
Peru and Bolivia. Northern populations are also 
broadly sympatric with two other species of the 
"Ophryoessoides group," S. aculeatus and S. fimbria- 
tus. However, in southern Peru S. prionotus is not 
known to be sympatric with other species of that 
group. I postulate that the higher vertebral crest in 
northern populations of S. prionotus functions as a 
species recognition signal in the multispecies assem- 
blages. A key to species of the "Ophryoessoides 
group" is provided and distributions of the species in 
Peru and Bolivia are summarized. 

Resumen. Se describe una nueva especie de lagar- 
tija iguanida, Stenocercus prionotus, del Peru Orien- 
tal y de Boli\aa. Se conoce la nueva especie desde el 
departamento de San Martin, Peni, hasta el norte de 
los departamentos La Paz y El Beni en Bolivia. La 
mayoria de las localidades se encuentran en las estri- 
baciones andinas y adyacentes tierras bajas. Stenocer- 
cus prionotus es similar a varias otras especies de 
Stenocercus con grandes escamas sobre el posterior 
de la cabeza, una fila amplia de supraoculares, y es- 
camas ventrales quilladas. Se refiere estas especies 
como el "grupo Ophryoessoides," sin implicar su 
monophyletismo. Basada en su posesion de un lobulo 
escamoso unico que oculta una porcion del bolsillo 
antehumeral, se considera la nueva especie cercana- 
mente relacionada a Stenocercus caducus (Cope), que 
se conoce desde Bolivia central hasta el norte de la 



Bull. Mus. Comp. ZooL, 157(3): 183-222, November, 2001 183 



184 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



Argentina y del Paraguay. Stenocercus prionotus se 
distingue de S. caduciis al tener una cresta vertebral 
mas prominente y un patron de alternativas barras 
claras y oscuras sobre la garganta (un patron rara- 
mente observado en S. caducus, que usualmente ti- 
ene manchas claras sobre la garganta). 

Las poblaciones de Stenocercus prionotus del norte 
del Peni tienen una cresta vertebral mas alta que la 
cresta en poblaciones del sur de Peru y Bolivia. Las 
poblaciones del norte de Peru tambien son amplia- 
mente simpatricas con dos otras especies del "gnipo 
Ophryoessoicles," S. aculeatus y S. finibriatus. Sin em- 
bargo, en el sur del Peni y Bolivia no se conocen lo- 
calidades donde se encuentra S. prionotus simpatrica 
con otras especies del giaipo. Postulo que la cresta ver- 
tebral mas alto en las poblaciones nortefias de S. prion- 
otus functiona como un serial para reconocimiento de 
especies en comunidades donde hay varias especies 
simpatricas. Se provee una clave para las especies del 
"grupo Ophnjoessoides" y se resumen las distribu- 
ciones de las especies Peruanas y Bolivianas. 

INTRODUCTION 

Stenocercus sensu Frost (1992), includ- 
ing the nominal genera Ophryoessoicles and 
Proctotretus, is a moderately diverse assem- 
blage of South American iguanid lizards 
(sensu Macey et al., 1997; Schulte et al., 
1998) with about 50 species currently rec- 
ognized. Most of the species are in the An- 
des and adjacent lowlands of Colombia, Ec- 
uador, and Pei"u, but a few are primarily 
Amazonian or have distributions in the 
physiographically diverse terrain south of 
the Amazon basin. Although new species of 
Stenocercus continue to be discovered in 
the field, others have been known from old 
collections and are only now being de- 
scribed (e.g., Cadle, 1991, 1998; Avila-Pi- 
res, 1995). In this category is a new species 
from the lowlands of eastern Peru and Bo- 
livia that has been referred erroneously to 
the names aculeatus O'Shaughnessy (1879) 
or caducus Cope (1862) in previous litera- 
ture, and associated with the genera Lei- 
ocephalus or Ophnjoessoides before the 
current understanding of these names 
came into use (see Etheridge, 1966; Frost, 
1992). Rodriguez and Cadle (1990) left the 
new species nameless in a checklist pend- 
ing resolution of its status. The new species 
is apparently closely related to Stenocercus 
caducus (Cope) and is described herein. 



MATERIALS AND METHODS 

Frost (1992; see especially footnote 5) 
and Cadle (1991) discussed reasons for re- 
ferring new species such as the one de- 
scribed here to Stenocercus Dumeril and 
Bibron sensu lato (including Ophnjoesso- 
ides Dumeril and Proctotretus Dumeril 
and Bibron). Externally, the new species is 
similar to those species that Fritts (1974) 
placed in Ophnjoessoides, that is, those 
species with keeled ventral scales, large 
posterior head scales (usually including 
well-differentiated interparietal, parietals, 
postparietals, and occipitals), and one 
moderately to greatly enlarged supraocular 
row. In addition to the new species de- 
scribed here, species included in the 
''Ophnjoessoides group" are aculeatus 
O'Shaughnessy, 1879; caducus Cope, 
1862; dunierilii Steindachner, 1867; ery- 
throgaster Hallowell, 1856; finibriatus Avi- 
la-Pires, 1995; huancahamhae Cadle, 
1991; iridescens Giinther, 1859; liniitaris 
Cadle, 1998; scapidaris Boulenger, 1901; 
tricristatus Dumeril, 1851; and two un- 
described species noted later in this paper 
under Key to Species of the "Ophryoesso- 
ides group" o/ Stenocercus. I use the term 
''Ophnjoessoides group" as a convenience 
to refer to this group of phenotypically 
similar species without implication as to its 
status as a monophyletic or nonmonophy- 
letic assemblage within Stenocercus. m 

General descriptive protocols follow Ca- 
dle (1991), who defined terminology of the 
scales, neck folds, and mite pockets used 
herein, based in part on Frost (1992). Bi- 
lateral scale counts (e.g., subdigitals) were 
made only on one side (the left, unless it 
was damaged), except for the holotype, for 
which both left and right counts were re- 
corded (1, r). A summary of selected scu- 
tellational and qualitative characters for 
the new species and similar species from 
eastern Peru and Bolivia is presented in 
Table 1. 

All measurements are in millimeters. 
The abbreviation SVL refers to the head- 
body length, from snout to vent. The con- 



New Species of Stenocercus from Peru and Bolivia • Cadle 



185 



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186 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 




Figure 1. Distribution of species of Stenocercus empfnasized in tfiis paper (western South America, Ecuador to Paraguay and 
northern Argentina). Open symbols for S. fimbhatus are literature records from Avila-Pires (1995); locality for S. caducus in 
northern Argentina is the southernmost locality in Argentina reported by Cei (1993). Otherwise, all records are based on spec- 
imens examined. Numbered localities are documented or suspected cases of sympatry referred to in the text and noted in the 
Appendix: 1, Pampa Hermosa; 2, Pampa Seca, Rio Mixiollo; 3, Tingo Maria; 4, Manu National Park. Upper left quadrant outlined 
with dotted line is the area shown in greater detail in Figure 2. 



figurations of neck folds and mite pockets 
vaiy considerably among species of Sten- 
ocercus and are useful in distinguishing 
species. The most important qualitative 
characteristics of these features used here- 
in are the following, which are discussed 
more fully by Cadle (1991): 

Neck and Body Folds and Crests. In 
contrast to many species of Stenocercus, 
neck folds are usually absent or weakly de- 



veloped in the "Opliryoessoides group." 
When present, they are better character- 
ized as crests rather than folds because 
they are usually indicated by strongly 
keeled rows of scales instead of actual 
folds of skin. The position of such crests 
corresponds to the position of folds seen 
in other species of Stenocercus, but only 
two are commonly seen in the "Ophryoes- 
soides group": an antehumeral crest, which 



New Species of Stenocercus from Peru and Bolivia • Cadle 



187 



is a more or less vertical row of strongly 
keeled scales immediately anterior to the 
forelimb insertion, and usually highlighted 
with white scales; and a supra- auricular 
crest, a strongly keeled horizontal row of 
scales extending from the posterior tem- 
poral region to the shoulder region. All 
species of the ^'Ophnjoessoides group" 
have a distinct vertebral crest formed by 
the strongly keeled, and often projecting, 
scales of the vertebral row. In addition, 
some of these species have a more or less 
prominent dorsolateral crest formed by a 
row of strongly keeled scales separating 
the flanks from the dorsum proper. The 
dorsolateral crest varies in length. In some 
species it is exceptionally long, extending 
from the proximal portion of the tail to the 
neck region, where it is confluent with the 
supra-auricular crest. In other species it is 
present only anteriorly or posteriorly. 

Posthiimeral (Axillary) and Postfemoral 
Mite Pockets. Type 1 — pocket absent; no 
skin modification. Type 2 — rudimentaiy 
pocket manifested by skin modification, 
such as bare skin, a series of wrinkles, or 
a shallow depression lined with scales dif- 
ferent from surrounding body scales. Type 
3 — similar to Type 2, but with an over- 
hanging fold of skin or a thickened border. 
Type 4 — a deep pocket, usually with a 
broad circular opening, whose depth is 
greater than half the diameter of its open- 
ing. Type 5 — a deep pocket with a narrow, 
slit-like opening and a depth greater than 
half the diameter of its opening. In two 
species discussed herein the posthumeral 
pocket is partially concealed by a scaly flap 
of skin, which I term a posthumeral or ax- 
illaiy flap. This structure is described more 
fully later. 

Angulate temporal scales are distinctly 
enlarged, keeled scales posterior to, and in 
line with, the superciliaiy scales. When 
present, they form a distinct border be- 
tween the posterior head scales and the 
lateral temporal scales, and they are mor- 
phologically distinguishable from these se- 
ries (Cadle, 1991:^6-7; see Fig. 4). Angu- 
late temporal scales are equivalent to su- 



pratemporals as used in some literature 
(e.g., Avila-Pires, 1995). In several species 
of Stenocercus the angulate temporals are 
not only keeled but they bear a projecting 
bladelike vane from the keels; in such cas- 
es I refer to the scales as "projecting." 

Coordinates for localities were obtained 
from the ornithological gazetteers of the 
Neotropics (Stephens and Traylor, 1983; 
Paynter, 1989, 1992, 1993, 1997), and 
from Lamas (1976), Morales and Mc- 
Diarmid (1996), Schulenberg and Awbrey 
(1997b), and Peruvian department maps 
produced by the Instituto Geografico Na- 
cional, Lima. I also consulted the on-line 
versions of the Peru and Bolivia gazetteers 
of the U.S. Board on Geographic Names 
at the GEOnet® Names Sei-ver: http:// 
164.214.2.59/gns/html/index.html. Brack- 
eted data in localities are inferences from 
these or other cited sources. Distributions 
of the new species and others emphasized 
in this paper are given in Figures 1 and 2. 
Institutional abbreviations are given at the 
beginning of the Appendix. 

DESCRIPTION OF A NEW SPECIES OF 
STENOCERCUS 

Stenocercus prionotus^ 
new species 
Figures 3-7, Figure 12; Table 1 

Liocephalus cadiicus (Cope, 1862): Boulenger (1898), 
specimen from "Barraca, Rio Madidi" [Boli\da] 
(BMNH 98.6.9.4) and probably two other northern 
BoHvian locahties discussed in the text. 

Ophnjoessoldes caducus (Cope, 1862): Fugler (1989: 
63), specimens from San Marcos Ranch, El Beni 
Department, Boli\da, including ROM 12815. 

Ophryoessoides acideatus (O'Shaughnessy, 1879): 
Fugler (1983, 1986, 1989), specimens from Tumi 
Chucua, El Beni Department, Bolix-ia (USNM par- 
atypes). 

Ophryoessoides sp.: Rodriguez and Cadle (1990), 
specimen from Cocha Cashu, Manu National Park, 
Madre de Dios Department, Peru (MCZ 150243). 



- Stenocercus pnonotus was recognized as new by 
R. Etheridge, P. E. Vanzolini, and E. E. Williams 
many years ago. Vanzolini and Williams applied the 
unpublished name Stenocercus dorsatus to labels of 
many specimens in various collections. 



188 Bulletin Museum of Comparative Zoologij, Vol. 157, No. 3 



7511^0° 




New Species of Stenocercus from Peru and Bolivia • Cadle 189 





Figure 3. Dorsal and ventral views of the holotype of Stenocercus prionotus (USNM 193683). Approximately xO.87. 



Figure 2. Northern Peru and Ecuador (see Fig. 1) showing distributions of species in the "Ophryoessoides group." Numbered 
localities are documented or suspected cases of sympatry referred to in the text and the Appendix (see Fig. 1 for names). Open 
circle at 06°S, 77°W is the type locality for Stenocercus aculeatus (Moyabamba, San Martin Department). All other localities are 
based on specimens examined; see Cadle (1991, 1998) for S. iridescens, S. limitaris, and S. huancabambae. The known 
distributions of all species are indicated by the localities plotted; however, the distribution of S. /r/descens continues farther north 
in western Ecuador than the area covered by the map. 



190 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



Holotype (Figs. 3—5). United States 
National Museum of Natural History 
(USNM) 193683 (field number WCS 
2421). PERU: Depto. Huanuco: Jardin 
Botanico de la Universidad Agraria de la 
Selva, Tingo Maria, vicinity of Rio Hual- 
laga, 670 m elevation [09°18'S, 75°59'W]. 
Adult male collected 29 June 1966 by 
Wade C. Sherbrooke. 

Faratypes from the Vicinity of the Type 
Locality. PERU: Depto. Huanuco: Ca. 2 
mi. by trail W. Tingo Maria, west bank of 
Rio Huallaga in vicinity of confluence with 
Rio Monzon, 670 m elevation (9 October 
1966, W. C. Sherbrooke), USNM 193685. 
Universidad Agraria de la Selva, Tingo 
Maria, Rio Huallaga, 670 m elevation (18 
August 1967, unknown collector for W. C. 
Sherbrooke), USNM 193686. Vicinity of 
Cueva de las Lechuzas, ca. 3 mi. SW Tingo 
Maria, Rio Monzon, ca. 700 m elevation 
(17 April 1968, Vito Yaringano for W. C. 
Sherbrooke), USNM 193687. Picuriacu, 
ca. 2 mi. NW Tingo Maria, Rio Huallaga 
(20 April 1968, W. C. Sherbrooke), USNM 
193688. 

Other Faratypes. PERU: Depto. 
Huanuco: Buena Vista, Valley of the 
Chimchao [ = Rio Chinchao] [approximate- 
ly 9°31'S, 75°52'W] (1-15 September 
1923, E[dmund] Heller), FMNH 5582- 
83. Hacienda Pampayacu [09°33'S, 
75°54'W] (17 July-16 August 1936, Dr. 
Snonge), MCZ 43758-59, 43761-62. 
Rio Llullapichis, 4—5 km upstream from 
Rio Pachitea, 200 m elevation [09°37'S, 
74°55'W] (Januaiy 1969, Hans W. Koep- 
cke), KU 179058. [Depto. Loreto]: 
E [astern] Peru, Pampa Hermosa, near 
mouth of Rio Cushabatay, Rio Ucayali Val- 
ley 500 ft. [152 m] [07°12'S, 75°17'W] 
(date unknown, H[ai-vey] Bassler), AMNH 
56760-64. Depto. Madre de Dios: Co- 
cha Cashu Biological Station, Manu Na- 
tional Park [11°51'S, 71°19'W] (July 1975, 
John W. Fitzpatrick), MCZ 150243. Ex- 
plorer's Inn, Tambopata Reserve, ca. 30 
km (straight line) SSW Puerto Maldonado, 
280 m [12°50'S, 69°17'W] (2 September- 
7 October 1983, native collectors), USNM 



247468-69, 247680; (23 May 1986, Vic- 
tor R. Morales), USNM 269022. Depto. 
Puno: Prov. Sandia: Tambopata, San Juan 
[del Oro], 1520 m or 5000 ft. [14°12'S, 
69°08'W]^ (22 November-20 December 
1950, Hilda H. Heller), FMNH 64788- 
92, 64794-811. [Depto. San Martin]: 
Juanjui [07°11'S, 76°45'W] (collector and 
date' unknown), MCZ 121233. Tarapoto, 
370 m [06°30'S, 76°25'W] (25 July 1984, 
Rainer Schulte), KU 212629. 

ROLIVIA: Depto. Reni: Provincia Va- 
cadiez, Tumi Chucua [176 m; 11°08'S, 
66°10'W] (23 October-18 November 
1981, Charles M. Fugler), USNM 
280246—51. Puerto Cruzeiro, San Marcos 
Ranch at confluence of Rios Isiboro and 
Ichoa [15°17'S, 65°45'W] (10 February 
1977, J. Lovisek), ROM 12815. [Depto. 
La Paz]: Barraca, Rio Madidi [12°35'S, 
67°02'W] (1893, Luigi Balzan), RMNH 
98.6.9.4. 

Distribution (Fig. 1). Stenocercus prion- 
otiis is known from the lowlands and An- 
dean foothills of eastern Peru and adjacent 
Bolivia (San Martin department, Peru, 
south to the Rio Beni in northern Bolivia). 
The known elevational range is 176—1,520 
m, but the highest recorded elevation is 
more than twice the elevation of the next 
lower one. Most localities in Peru are ad- 
jacent to or near the Andean foothills, and 
several localities in La Paz and El Beni 
departments, Bolivia, are unconfirmed. No 
specimens are known from a broad geo- 
graphic hiatus between central and south- 
ern Peru (Fig. 1). See Distribution Fat- 
terns in Stenocercus prionotus for further 
discussion. 

Etymology. The epithet prionotus is de- 



^ Hilda Hellers notes on this collection in the Karl 
P. Schmidt archives of the FMNH describe San Juan 
as the site of an agricultural station on "the left side 
of die Rio Tambopata at 5000 feet." With some hes- 
itancy I identify Heller's locality as the town kno\\'n 
as San Juan del Oro, which is on the left bank of the 
Rio Tambopata at approximately the elevation given 
by Heller. I have located only two other places named 
San Juan in Puno Department, but neither is on the 
Rio Tambopata. 



New Species of Stenocercus from Peru and Bolivia • Cadle 



191 



rived from the Greek adjective prionotos 
meaning jagged or serrate. The name re- 
fers to the strongly serrate vertebral crest 
of Stenocercus prionotus, which is the 
most obvious character distinguishing this 
species from its apparent closest relative, 
S. caduciis. 

Data on the Holotype. Adult male, hem- 
ipenes partially everted. SVL, 83 mm. Tail 
length, 201 mm. Total length, 284 mm. 
Tail/total length, 0.71. Vertebral scales be- 
tween the occipital and the posterior mar- 
gin of the hind limb, 31. Midbody scales, 
42. Gular scales between the ears, 16. In- 
ternasals, 6. Subdigital scales on fourth fin- 
gers and toes, respectively, 18—18, 25—25. 
Color pattern well presei-ved: top of head 
brown with narrow dark brown interorbital 
bar extending laterally onto supraoculars; 
dorsum brown with narrow blackish chev- 
rons middorsally (1 on neck, 1 above fore- 
limbs, 2 others anterior to midbody; pos- 
terior chevrons poorly defined); dark 
brown scapular blotch bordered anteriorly 
by white antehumeral stripe; ill-defined 
grayish dorsolateral streaks between ear 
and anterior body; throat grayish with 
poorly defined oblique light grayish 
stripes; venter brown without distinct pat- 
tern. 

Definition. A species of Stenocercus 
characterized by the following features: (1) 
Dorsal head scales subimbricate and 
strongly keeled to multicarinate; temporals 
keeled, imbricate or subimbricate. (2) Pos- 
terior head scales larger than anterior 
ones, with distinct interparietal, a pair of 
parietals, a pair of postparietals, and a 
large median occipital (often surrounded 
by several small irregular scales). (3) In- 
ternasals usually 7, but pattern irregular 
and may be 5 or 6. (4) One row of supra- 
oculars distinctly enlarged. (5) One canthal 
on each side between the superciliaries 
and the lateralmost internasal. (6) A pair 
of strongly keeled angulate temporals in 
line on each side (rarely, 3 angulate tem- 
porals are present), each with a low pro- 
jecting blade; partially or completely sep- 
arated from enlarged posterior head scales 



by a single row of small scales. (7) Anterior 
and posterior gular scales strongly keeled. 
(8) Parietal eye distinct. (9) Neck folds ab- 
sent; a vertical, strongly keeled row of 
scales in the antehumeral region and oc- 
casionally a much less distinct raised series 
in the supra-auricular region. (10) Dorsal 
and ventral body scales imbricate, mucro- 
nate, strongly keeled; dorsal scales at mid- 
body 36—48. (11) Vertebral row continu- 
ous, bearing a strongly projecting serrate 
crest in adults; a dorsolateral crest present 
on posterior body and the base of the tail. 

(12) Deep posthumeral pocket (Type 4) 
partially concealed by a scaly posthumeral 
flap originating on its anteroventral bor- 
der; postfemoral pocket absent (Type 1). 

(13) Scales of posterior thigh imbricate, 
keeled. (14) Tail strongly compressed in 
adults, anteriorly with low vertebral and 
dorsolateral crests continuous with those 
of the body. (15) Dorsal coloration of 
males in preservative (Figs. 3, 6) brown 
with or without distinct chevrons; a dis- 
tinct white vertical antehumeral stripe ex- 
tending ventrally to the proximal ventral 
surface of forelimb; a large dark scapular 
blotch; in well-presei"ved specimens the 
throat bears oblique alternating dark and 
light stripes (see Description); females 
similar but pattern elements often more 
subdued. 

DIAGNOSIS AND COMPARISONS 

In having enlarged posterior head 
scales, an enlarged row of supraoculars, 
and strongly keeled ventral scales, Steno- 
cercus prionotus is like other species in the 
"Ophryoessoides group" of Stenocercus. 
These are the species most likely to be 
confused with S. prionotus. Five other de- 
scribed species of the ''Ophryoessoides 
group" occur in eastern Peru or Bolivia: S. 
aculeatus (O'Shaughnessey, 1879); S. cad- 
ucus (Cope, 1862); S. finibriatus Avila-Pi- 
res, 1995; S. huancabanibae Cadle, 1991; 
and S. scapularis (Boulenger, 1901). An 
undescribed species occurs in the Rio 
Maraiion valley of eastern Peru (see Key 
to Species of the "Ophryoessoides Group" 



192 Bulletin Museum of Comparative Zoologtj, Vol. 157, No. 3 



of Stenocercus). Stenocercus prionotus 
and S. caducus are unique among known 
species of Stenocercus (perhaps unique 
within iguanids) in having deep posthu- 
meral mite pockets (Type 4) that are par- 
tially concealed anteroventrally by a scaly 
flap, which may be termed a posthumeral 
or axillary flap (Fig. 5). Stenocercus prion- 
otus and S. caducus are compared in 
greater detail below, but S. pmonotus is 
distinguished from S. caducus (character- 
istics in parentheses) by: (1) a strongly pro- 
jecting, serrate vertebral crest (low and 
scarcely projecting); (2) 2 (usually) or 3 en- 
larged, strongly keeled, projecting angu- 
late temporal scales on each side (scales 
not greatly enlarged, less projecting); and 
(3) a gular pattern consisting, when evi- 
dent, of oblique alternating dark and light 
lines or bars, or oblique light lines on a 
dark ground color (usually light spots on a 
darker ground color, unicolor, or [rarely] a 
pattern similar to that of S. prionotus). 

Readily determined characters distin- 
guishing Stenocercus prionotus from the 
other four species of the "Ophryoessoides 
group" known from eastern PeiTi and Bo- 
livia include the extent of keeling on dorsal 
head and body scales, relative develop- 
ment of the postfemoral pockets, and the 
number of midbody scale rows (Table 1 
and key presented later herein). Stenocer- 
cus fimbriatus and S. aculeatus are known 
to be sympatric with S. prionotus at several 
localities in eastern Peru. In addition to 
having a posthumeral flap (absent in S. 
fimbriatus and S. acideatus), S. prionotus 
is distinguished from S. fimbriatus (char- 
acteristics in parentheses; see Avila-Pires, 
1995) in having strongly keeled dorsal 
scales in adults (smooth or weakly keeled), 
a dorsolateral crest prominent only on the 
posterior body (prominent anteriorly and 
continuous with antehumeral and supra- 
auricular folds or crests), and in lacking 
"fimbriate" scales on the posterior distal 
portion of the thigh (present). Stenocercus 
prionotus is distinguished from S. aculea- 
tus (characteristics in parendieses) in lack- 
ing a postfemoral pocket (moderate to 



deep); in having strongly keeled, often 
multicarinate, head scales (smooth or 
weakly striated in adults, wrinkled in ju- 
veniles); 5—7 internasals (4—5); and only 
moderately enlarged supraoculars, usually 
5—6 supraoculars across the widest part of 
the orbit (greatly enlarged, usually 4 across 
the orbit). 

Stenocercus prionotus differs from S. 
scapularis (characteristics in parentheses) 
in lacking squarish or rectangular project- 
ing superciliaiy scales (present) and in 
having fewer than 50 midbody dorsal scale 
rows (59—70 rows). Stenocercus prionotus 
differs from S. huancabambae (characters 
in parentheses) in lacking a postfemoral 
pocket (deep, Type 5) and in having prom- 
inent dorsolateral crests on the posterior 
body (weak, restricted to anterior body 
when present). 

Two species of Stenocercus from west- 
ern Ecuador and Peru, S. iridescens and 
S. limitaris, have enlarged posterior head 
scales and supraoculars. In contrast to 
Stenocercus prionotus, S. iridescens has 
smooth head plates, 2 canthals, a poorly 
developed posthumeral pocket (Type 1 or 
2), and lacks keeled angulate temporals 
and dorsolateral crests on the body (see 
Cadle, 1991, fig. 10). Stenocercus limitaris 
has a deep postfemoral pocket (Type 5), 2 
canthals, a single strongly keeled (but non- 
projecting) angulate temporal, and lacks 
dorsolateral crests. 

Other non-Peruvian species of the 
"Ophryoe.ssoides group" can be distin- 
guished from Stenocercus prionotus by 
features in the key presented later and 
other superficial characters, such as the 
presence of 2 canthals and 4 internasals 
(usually) in S. erythrogaster (1 and 5—7 in 
S. prionotus), and enlarged, projecting py- 
ramidal or conical postsuperciliary scales 
in S. dumerilii and S. tricristatus (Avila- 
Pires, 1995). 

Other species of Stenocercus are distin- 
guished from S. prionotus by a combina- 
tion of features such as smaller head 
plates, smooth ventrals, and absence of 
dorsolateral crests. Most species of Steno- 



New Species of Stenocercus from Peru and Bolivia • Cadle 



193 



cercus except the "Ophnjoessoides group" 
have smooth or weakly keeled head plates. 
Other characters, such as the number of 
dorsal scale rows, morphology of the pos- 
thumeral and postfemoral mite pockets, 
extent of sexual dimoi*phisin, and degree 
of differentiation of the vertebral scale row 
and crest also aid in distinguishing the spe- 
cies (see descriptions and discussion in 
Fritts, 1974; Cadle, 1991, 1998). 

Apart from Stenocercus fimbriatus, S. 
scapularis, and S. acideatus, two other spe- 
cies of Stenocercus are known from local- 
ities close to or synipatric with known pop- 
ulations of S. prionotus: S. crassicaudatus 
and S. roseiventris. These species are dis- 
tinguished from S. prionotus by lacking 
prominent serrate vertebral crests (low 
crests may be present), having smooth 
ventral scales, and having prominently spi- 
nose tails with the spines arranged in dis- 
tinct whorls. 

DESCRIPTION 

Head (Fig. 4). Dorsal head scales sub- 
imbricate (a tendency to be more juxta- 
posed posteriorly); strongly keeled to mul- 
ticarinate or wrinkled. Rostral in contact 
with first supralabial, first lorilabials, and a 
series of postrostrals. Usually 7 elongate, 
strongly keeled internasals between the 
nasals dorsally; however, the anterior dor- 
sal head scales are very irregular and oc- 
casionally only 5 or 6 internasals are pre- 
sent. One canthal scale between the an- 
terior superciliary and the lateralmost in- 
ternasal, separated from the nasals by tiny 
postnasals. Canthus very strongly angled. 
Nostril in posterior portion of an elongate 
nasal scale, which may contact the rostral 
scale anteriorly or be separated from it by 
small postrostrals. Four or 5 strongly over- 
lapping, elongate anterior superciliaries 
followed by 2 or 3 shorter posterior su- 
perciliaries slightly overlapping in the re- 
verse direction (but more or less in a 
straight line). One supraocular row inod- 
erately enlarged, 2 mediocentral scales 
much larger than the others. Five or 6 
scales across the supraocular area at its 



widest part. Interparietal distinct and elon- 
gate, diamond-shaped or pentagonal (apex 
posteriorly). Parietal eye visible. A pair of 
parietals in contact behind the inteqDari- 
etal, flanked posterolaterally by a postpar- 
ietal on each side. Postparietals separated 
medially by a single median transversely 
elongate occipital; occasionally 1 or 2 small 
scales are intercalated at the juncture of 
the parietal, postparietal, and/or occipital 
(e.g.. Fig. 4). 

Lateral temporal scales strongly keeled, 
imbricate to subimbricate; separated from 
posterior dorsal head scales on each side 
by 2 (occasionally 3) elongate, strongly 
keeled angulate temporal scales bearing a 
low projecting vane. Keels of adjacent an- 
gulate teinporals aligned. Posterior angu- 
late temporals separated from postparie- 
tals by 2 or 3 small scales in a longitudinal 
row. Anterior angulate temporals may con- 
tact postparietal and one other larger pos- 
terior head scale or be separated from 
them by small scales. Anterior border of 
ear weakly denticulated; posterior border 
rounded, bordered with keeled imbricate 
scales. 

Anterior and posterior gulars strongly 
keeled. Mental smooth, in contact with 
first pair of postmentals and first pair of 
infralabials. Enlarged postmentals 3 or 4 
on each side, only the first pair in contact 
medially. 

Neck and Body. Dorsal and lateral 
scales of neck and body imbricate, mucro- 
nate, strongly keeled. Vertebral row pro- 
duced into a prominent projecting serrate 
crest in adults of both sexes that is contin- 
uous from the nuchal region to the base 
of the tail, gradually disappearing on the 
anterior Va to V2 of tail. Dorsolateral crest 
(a raised, strongly keeled row of scales) on 
posterior % of body, continuing onto base 
of tail. The dorsolateral crest occasionally 
appears very indistinctly farther anteriorly 
on the body, but only on the posterior 
body does it shaiply delimit the dorsolat- 
eral (paradorsal) scales from the flank 
scales. Three rows of scales between dor- 
solateral and vertebral crests at anterior 



194 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 




Figure 4. Head scales of the holotype of Stenocercus prionotus (USNM 193683) in dorsal and lateral views. Scale bars = 3 
mm. To facilitate coordination with the text, the following scales are indicated on one side (interparietal and occipital are median 
scales); A, angulate temporal; I, interparietal; O, occipital; P, parietal; PP, postparietal. 



New Species of Stenocercus from Peru and Bolivia • Caclle 



195 



edge of dorsolateral crest (i.e., just anterior 
to the pelvic region), 2 rows posteriorly 
(dorsal to hindlimbs); scales between the 
crests strongly imbricate and keeled only 
on posterior part of scale. Flank scales 
mostly fully keeled (sometimes only the 
posterior part of each scale), imbricate, 
mucronate, slightly smaller than dorsolat- 
eral scales. Ventral body scales mucronate, 
strongly keeled (keels running the length 
of the scales). Ventrals approximately the 
same size as the dorsolateral scales, larger 
than flank scales. 

Neck Folds. Distinct neck folds absent. 
Poorly developed antehumeral crest pre- 
sent. 

Tail. Tail strongly compressed, anteriorly 
bearing low projecting vertebral and dor- 
solateral crests continuous with those of 
the body. Dorsal scales moderately keeled, 
ventral scales strongly keeled. 

Limbs. Dorsal and ventral scales of fore- 
limbs, hindlimbs, and posterior thigh 
strongly keeled, unicarinate, mucronate; 
some scales of the shank larger than any 
thigh scales. Supradigitals and subdigitals 
unicarinate. Palmar scales strongly multi- 
carinate. Plantar scales strongly unicari- 
nate. 

Posfhiimeral and Postfemoral Mite 
Pockets. Posthumeral mite pocket a deep 
cavity (Type 4) with a prominent axillaiy 
flap concealing the anteroventral aspect 
(Fig. 5). Postfemoral pocket absent (Type 

The flap associated with the posthumer- 
al pocket projects from the anteroventral 
and ventral edges of the pocket. Antero- 
dorsally, a similar but much smaller flap is 
present in some specimens (e.g.. Fig. 5). 
The posthumeral flap consists of a fleshy 
ridge covered anteriorly and posteriorly 
(or externally and internally when the flap 
is lying flat against the body) by keeled im- 
bricate scales. Externally, usually 3 or 4 
larger scales cover the flap ventrally and a 
series of much smaller scales is present 
dorsally. One or 2 of the larger scales are 
sometimes highlighted with white. When 
appressed against the body (i.e., against 




Figure 5. Posthumeral (axillary) flap of Stenocercus priono- 
tus (USNM 193683, holotype). Anterior to the right. The broad 
oval on the right is the deflected forelimb and the posthumeral 
mite pocket is the heavily stippled cavity deep to the flap. The 
posterior border of the axillary flap is marked by the scales 
with heavily outlined posterior borders and it extends anteriorly 
to the ventral part of the forelimb. A smaller dorsal flap is also 
present in this specimen (small patch of heavily outlined scales 
on anterodorsal edge of the pocket; see text). Scale bar = 1 
mm. 



the opening of the posthumeral pocket) 
the flap conceals approximately the ventral 
Vs to Vi of the vertical dimension of the 
pocket. The flap is equally prominent in 
adults of both sexes and is proportionally 
as well developed in subadults (including 
hatchlings) as in adults. 

Size and Propoi~tions. Largest inale 
(USNM 193683) 89 mm SVL, 323 mm to- 
tal length (sample size of males with SVL 
>70 mm = 13). Largest female (KU 
212629) 93 mm SVL, 329 mm total length 
(sample size of females with SVL >70 mm 
= 16). Tail relatively long, 69-74% of total 
length in adults (67-71% in juveniles). 

Coloration and Pattern of Adult Males 
in Life. The following color descriptions 
are paraphrased from the field notes of 
Wade C. Sherbrooke. USNM 193683 (ho- 
lotype): 

Lower half of side between limbs is lavender- 
brown. This color extends from both sides across 
the belly approximately Vs of the way on each side, 
leaving a tan-brown central strip down the belly. 
General base color of the body is brown, darkest 



196 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 








Figure 6. Lateral view of Stenocercus prionotus from northern Peru (USNM 193685; male, snout-vent length 76 mm). Note 
the high vertebral crest and the following pattern elements: dark subocular bar, dark blotches on the dorsal and lateral surfaces 
of the neck, pale antehumeral bar, indistinct pale dorsolateral stripe, and indistinct oblique bars on the trunk. 



near the hind legs and tan just in from of the fore- 
legs. A distinct white line runs dorsally from the 
top of each foreleg to three quarters of the way to 
the dorsal crest; it runs through a large black patch 
just above the forelimb. Head markings consist of 
a dark brown line i-unning between the eyes on die 
top of die head; this continues through the eye to 
broaden slightly at the rear portion of the jaw. The 
gular area is streaked by several light cream lines. 
Very slight lavender tinge to all of body behind 
forelimbs. [Sketch in notes shows black middorsal 
patches that don't extend to flanks]. 

USNM 193685 (WCS 2543): 

This specimen closely resembles [USNM 193683] 
in color, with one exception. There are several 
green spots on . . . the right dorsal surface between 
the limbs and on the dorsal portions of the tail 
behind the hind limbs and the dorsal tibio-fibula 
portion of the right hind leg. 

Coloration of Adult Females in Life. Un- 
known. 

Coloration in Preservative (Figs. 3, 6, 7). 
Specimens of Stenocercus prionotus vary 
greatly in coloration due mainly to varia- 
tion in initial preservation and the length 
of time in presei-vative. Poorly preseived 
specimens may be more or less uniform 
brown all over, although obscure pattern 
eleiTients are usually present. Well pre- 
served specimens are brown dorsally with 
darker brown to blackish chevrons mid- 
dorsally. One chevron dorsal to each pair 
of limbs and one on the neck are usually 
evident, and these are darker than others 
that iTiay be present. Three to five mid- 
dorsal chevrons are between the limbs. 



Dark spots or an additional chevron are 
often present on the dorsal neck and usu- 
ally on the base of the tail. The light an- 
tehumeral/humeral line is universally pre- 
sent and evidence of the dark shoulder 
patch is usually present (often veiy prom- 
inent). Flanks usually unicolor and some- 
what darker than the dorsum between the 
dorsolateral crests; however, soixie speci- 
mens (e.g., KU 179058, USNM 280246) 
have distinct dirty white vertical bars or 
chevrons on the flanks (five between the 
limbs), and such bars appear occasionally, 
but more obscurely in other specimens 
(see Fig. 6). The dorsolateral crest is often 
highlighted for a variable length with a dis- 
tinct or indistinct light line, giving the im- 
pression of a light dorsolateral stripe. 
Forelimbs more or less unicolor brown or 
with obscure pattern; hindlimbs brown 
with darker brown bands. Dark subocular 
bar distinct. Top of head often with an ob- 
scure or distinct dark brown interocular 
bar. Oblique bars on throat (Fig. 7) often 
visible but throat may be unicolor or have 
an obscure pattern. Venter of most speci- 
mens unicolor, dirty white, gray, or beige; 
however, some specimens (e.g., USNM 
280246) have a series of irregular longi- 
tudinal dark brown streaks. 

Scale Counts and Qualitative Features 
(Table 1). Stenocercus prionotus has rela- 
tively low midbody, vertebral, and gular 
scale counts. The scales are relatively large 
and strongly keeled over most of the body. 



New Species of Stenocercus from Peru and Bolivia • Cadle 



197 





Figure 7. Gular patterns in Stenocercus prionotus. Top, typ- 
ical throat pattern (oblique view) consisting of light and dark 
stripes that extend medially to the midline (USNM 193687). 
Bottom, specimen in which the ground color is lighter and 
therefore the contrasting pale stripes are less distinct (USNM 
193685). 



Sexual Dimorphism. Stenocercus prion- 
otus does not exhibit strong sexual dimor- 
phism. Males and females attain approxi- 
mately the same size and have the same 
general pattern, but whether the colora- 
tion in life reported above for adult males 
pertains to females as well is unknown. 
The vertebral crest is only slightly more 
developed in males than in females of the 
same population but this character shows 
strong clinal variation (northern popula- 
tions with higher crests; further discussed 
below). Other characters that sometimes 
vary between the sexes in Stenocercus 
show little variation in S. priontotus. Nei- 



ther standard meristic counts (Table 1) nor 
the relative development of the posthu- 
meral and postfemoral pockets (Types 4 
and 1, respectively, in adults of both sexes 
and in subadults) show obvious sexual di- 
morphism. 

DISTRIBUTION PATTERNS IN 
STENOCERCUS PRIONOTUS 

Absence of Stenocercus prionotus from 
Lowland Localities in Eastern Fern. Sten- 
ocercus prionotus is widespread in the 
lowlands along the Andean front from 
northern Peru to northern Bolivia (Fig. 1). 
However, all Peruvian localities are close 
to the Andean foothills and south of the 
broad extension of the Cordillera Oriental 
separating the great bend of the Rio Mar- 
aflon from upper reaches of the Rio Hual- 
laga (Fig. 2). The absence of specimens in 
comprehensive collections from the Iqui- 
tos region (Dixon and Soini, 1986), Balta 
(Ucayali Department; specimens at 
LSUMNS and University of Arizona), and 
Cuzco Amazonico (Madre de Dios De- 
partment; Duellman and Salas, 1991) sug- 
gest that S. prionotus may be absent from 
the lowlands distant from the Andean foot- 
hills, at least in Peru. Similarly, collections 
from northern Loreto Department (Duell- 
man and Mendelson, 1995) and northern 
Amazonas Department (J. E. Cadle and R. 
W. McDiarmid, unpublished data from the 
Rio Cenepa and Rio Santiago) suggest that 
S. prionotus does not occur north of the 
Rio Marafion. 

However, these sites have been sampled 
unevenly. For example, species accumula- 
tion curves for lizards at Cuzco Amazonico 
(Duellman and Koechlin, 1991) reached 
an asymptote after about 15 person-weeks 
of effort, whereas only 5 person-weeks 
were expended in northern Loreto and the 
species accumulation curve for the total 
herpetofauna showed no asymptote 
(Duellman and Mendelson, 1995; data not 
presented for lizards only). Quantitative 
data are not available for the other sites, 
but large collections are available for the 
Iquitos region, including more than 1,000 



198 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



Table 2. Patterns of presumed sympatry of Stenocercus in eastern Peru. Only species of the 
"Ophryoessoides group" and the superficially similar species, S. roseiventris, are listed. 1 Locali- 
ties ARE listed roughly NORTH TO SOUTH; FOR PRECISE LOCALITIES SEE THE APPENDIX AND LOCALITIES 

FOR THE TYPE SERIES OF S. PRIONOTUS.. 



Iquitos San Martin/ Pampa 
Region VV. Loreto- Hermosa 



Rio 
Mishollo 



Tingo 
Maria 



Balta 



Cuzco 
Amazonico 



Explorer's 
Inn 



S. prionotus 




X 


X 




X 


X 


S. aculeatus 




X 




X 






S. fimbriatus 


X 




X 


X 


X 


X 


S. roseiventris 


X 










X 



^ The only other species of Stenocercus known from primarily the lowlands and lower Andean foothills in 
this region is S. crassicaiidatus, which lacks a projecting vertebral crest and has a very spiny tail. 

^ The distributions of S. prionotus and S. aculeatus overlap both altitudinally and latitudinally in northern 
Peru but they have not been taken together at the same locality. See Figure 2 for localities. 

^ In Manu National Park, S. fimbriattis and S. roseiventris are sympatric at Pakdtza, whereas S. prionotus is 
known only from Cocha Cashu. 



lizards obtained by Dixon and Soini (1986) 
and additional collections froin the region 
made by Harvey Bassler and deposited in 
the AMNH. Similarly large collections re- 
sulted from the efforts of Cadle and 
McDiarmid in Amazonas (specimens in 
the MVZ and USNM). Thus, barring ar- 
tifacts introduced by the difficulties of col- 
lecting cryptic rainforest lizards, S. prion- 
otus seeiTis to be absent froixi these sites. 

Documenting and explaining patterns of 
absence is always difficult, but the appar- 
ent absence of Stenocercus prionotus froin 
the lowlands distant from the Andes in 
Peru is not due to failure to collect Sten- 
ocercus at these localities because at least 
one other species of Stenocercus is known 
from each (Table 2). However, no lowland 
locality is known in which inore than two 
species of the "Ophryoessoides group" of 
Stenocercus are syinpatric. Of interest in 
this connection is that S. prionotus has not 
been taken in the upper reaches of the Rio 
Perene and its tributaries (Junin Depart- 
ment), although it is known from north 
and south of that region. Numerous spec- 
iinens of Stenocercus of at least five spe- 
cies (S. hoettgeri, S. crassicaudatus, S. for- 
mosus, S. scapularis, and S. variabilis'^) 



have been collected along an elevational 



^ Based on examination of two of the three synty- 
pes of Stenocercus variabilis Boulenger (BMNH 
1946.8.11.89, 1946.8.11.91) I concur with Fritts 
(1974: 66) that Boulenger (1901: 553) erred in as- 



cribing these specimens to "Palca, Bolixda" rather 
than Palca (Junin Department), Peru. One specimen 
I collected near Palca, Peru (MCZ 178166) is nearly 
identical with BMNH 1946.8.11.91 in scale counts, 
pattern, and qualitative characters (both of these 
specimens differ considerably from BMNH 
1946.8.11.89 in color pattern). However, the attri- 
bution of the specimens to Peru is not without some 
equivocation. The types were collected by P. O. Si- 
mons (see Cadle, 1998: footnote 6), who collected in 
the vicinity of Palca, Peru, in March and April, 1900, 
although that specific locality is not listed in his pub- 
lished itinerary (Chubb, 1919). However, Simons's 
itineraiy places him at "Palca, 18 miles E of La Paz" 
[Bolivia] on 9 November 1900 (Chubb, 1919: 5). Si- 
mons's field tags attached to the syntypes record sim- 
ply "Palca 3000 m," which is close to the elevation of 
Palca, Peru (2,740 m), but not that in Bolivia (4,600 
m). Adding to the confusion are entries in the 
BMNH registries for the syntypes, concerning which 
Colin J. McCarthy provided the following comments 
via e-mail: 

Boulenger originally wrote "Palca Peru 3000m" but 
later struck through Peru and wrote "Bolivia." 
There are two sheets of notes (presumably from 
Simons) stuck in at this page of the register about 
the localities in the batch. With regard to Palca he 
has written "Palca, just S. of La Paz, Bolivia." I 
assume it was that information that caused Boulen- 
ger to alter his original entry! 

Thus, the confusion may have originated with the 
note that Simons provided subsequent to cataloguing 
of the collection at the BMNH. In any case, no spec- 
imens resembling S. variabilis are definitely known 
from Bolivia and the only species of the genus defi- 
nitely known to occur above 4,000 m is Stenocercus 
chrysopt/gus from northern Peru. 



New Species of Stenocercus from Peru and Bolivia • Cadle 



199 



transect along this well-traveled route 
(Fritts, 1974; Cadle, 1991 and unpublished 
data). Thus, analysis of circumstantial dis- 
tributional data suggests that the distribu- 
tion of S. prionotus in the lowlands of east- 
ern Peru may be influenced by the num- 
ber of sympatric species of Stenocercus. At 
all localities from which S. prionotus has 
been taken, only one other species of Sten- 
ocercus is known (Table 2). These patterns 
of sympatry are discussed in the next sec- 
tion with reference to patterns of geo- 
graphic variation in S. prionotus. 

In contrast, the apparent restriction of 
Stenocercus prionotus to the Andean foot- 
hills and immediately adjacent lowlands in 
Peru does not seem related to the distri- 
bution of any major habitat type or phys- 
iographic region. Most of the extensively 
sampled localities (e.g., Iquitos and Cuzco 
Amazonico) include a variety of lowland 
habitats characteristic of western Amazon- 
ia. Stenocercus prionotus is known from 
both floodplain forests (Cocha Cashu) and 
more upland forests on river terraces in 
southeastern Peru (Explorer's Inn); Foster 
(1990) and Dallmeier et al. (1996) de- 
scribed these floristic communities in 
southeastern Peru. Thus, the apparent ab- 
sence of S. prionotus at the localities dis- 
cussed above is not due to some simple 
relation to local habitat availability. For ex- 
ample, it is unclear why S. prionotus was 
not obtained at Pakitza (Morales and 
McDiarmid, 1996), even though it occurs 
at nearby Cocha Cashu. On a broader geo- 
graphic scale, the restriction of S. priono- 
tus to lowlands and foothills adjacent to 
the Andes may be related to the present 
or historical influence of the Andes on the 
climate and vegetation (rainfall, tempera- 
ture, and major soil types) of neighboring 
regions. 

A curious and unexplained hiatus in the 
distribution of Stenocercus prionotus oc- 
curs between the vicinity of Tingo Maria— 
Rio Llullapichis (Huanuco Department) 
and Cocha Cashu in Manu National Park 
(Madre de Dios Department), a gap of 
some 600 km that includes the entire up- 



per reaches of the Rio Ucayali-Urubam- 
ba— Ene system. Additionally, populations 
north and south of this gap differ in some 
qualitative and quantitative characters (see 
Patterns of Sympatry and Geographic Var- 
iation in Stenocercus prionotus, and the 
Need for Additional Fieldivork). Scattered 
collections (e.g., maps in Fritts, 1974; Avi- 
la-Pires, 1995), but no comprehensive her- 
petofaunal surveys or collections, are avail- 
able from this vast region. Thus, whether 
the geographic hiatus is real or a sampling 
artifact cannot be discerned. Efforts to re- 
solve this issue need to be made. 

Unconfirmed Bolivian Localities for 
Stenocercus prionotus. Because of the 
previous confusion of Stenocercus priono- 
tus wath S. caducus, specimens perhaps re- 
ferable to S. prionotus from several local- 
ities in Bolivia are unconfirmed. Boulen- 
ger (1898) reported specimens of "Lioce- 
phalus caducus. Cope" in the Museo 
Civico Storia Naturale Giacomo Doria in 
Genoa collected by Luigi Balzan from four 
localities (Balzan, 1931). I have not at- 
tempted to verify the existence of these 
specimens. However, one of Balzan s spec- 
imens from "Barraca, Rio Madidi" was ex- 
changed to the BMNH (now BMNH 
98.6.94), and is confirmed as S. prionotus. 
The "Leiocephalus caducus" specimens 
from the other three Balzan localities are 
outside the known distribution of S. cad- 
tictis but are close to other known localities 
for S. prionotus in northern Bolivia (Fig. 
1). I suspect these are S. prionotus based 
on geographic location. The localities are, 
as listed by Boulenger (1898) (see Fig. 1), 

(1) "Coroico and Chulumani, Prov. Yun- 
gas, 1,600 metres alt." [La Paz Depart- 
ment; Coroico, 1,725 m, 16°10'S, 67°44'W; 
Chulumani, 1,905 m, 16°24'S, 67°31'W]. 

(2) "Reyes, right bank of Rio Beni" [El 
Beni Department, 232 m; 14°19'S, 
67°23'W]. (3) "Missiones [sic] Mosetenes" 
[approximately 15°31'S, 67°25'W].5 These 



^ The Moseten Indians inhabited upper reaches of 
the Rio Beni and its tributaries in the Andean foot- 
hills of the present department of La Paz (Metraux, 



200 



Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



localities would not be unusual for S. 
prionotus, although the first two localities 
are the highest elevations recorded for the 
species (the species occurs at 1500 m in 
nearby Puno Department, Peru). All are 
in the upper reaches of the Rio Beni, 
whereas the two confirmed Bolivian local- 
ities for S. prionotus are farther north in 
the same drainage. 

Burt and Burt (1931: 273) listed two 
specimens of ''Leiocephalus [ = Stenocer- 
cus] scapularis" (AMNH 22450, 22532) 
from Rurrenabaque, Bolivia [El 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., Februaiy 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 SYMPATHY AND 
GEOGRAPHIC VARIATION IN 
STENOCERCUS PRIONOTUS, AND THE 
NEED FOR ADDITIONAL FIELDWORK 

Stenocercus prionotus varies geographi- 
cally in several characters, most notably in 
the height of the vertebral crest. Because 



1942). According to Metraux (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. 



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 
"Ophiyoessoides 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 Huanuco and 
Puno departments (Peru) (Fig. 1; Appen- 
dix), and it seems to be elevationally par- 
apatric to S. prionotus in northern Peru 
(Huanuco 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. scaptdaris 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 



201 



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. finihriatus 
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 
"Ophnjoessoides group." However, these 
species have been taken together at only 
one locality. Explorer's Inn. 

To summarize these patterns of sympat- 
ly, 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 Huanuco and 
Loreto departments. No species of the 
"Ophrijoessoides 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 fiinbriafiis from Pak- 
itza (Appendix) extends the known range for tliis spe- 
cies south by approximately 200 km from Balta in 
Ucayah Department, Peru (Avila-Pires, 1995). 



er scales, of the crest in southern popula- 
tions. None of these differences in scale 
counts is statistically significant. 

The height of the vertebral crest shows 
strong clinal variation in Stenocercus 
prionotus. Populations from northern and 
central Peru have higher crests than those 
from southern Peru and Bolivia (Fig. 10). 
However, the absence of specimens be- 
tween Huanuco 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 dimoiphic 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 intei"vening 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 
Endler s (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- 



202 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



48 




• 
















47 




• 
















46 




















45 




• 


• 














44- 




•• 






• 






■ 




43 


• • 


• • 






• 










42 




••• 


• • 




•••• 




■ 


■ 


■ 


M 

0) 

5 41- 










•••• 










(0 




















■o 40- 
o 

Si 




• 


• • 


• 


• ••• 




■■ 




■ 




















i 39 




• 


• 


• 


• ••• 


■ 




■ 




38- 


•• 








• ••• 




■ ■■■ 




■ 


37- 






• 


• 






■■■■■ 
■ ■■■ 






36- 








•• 


• 




■■■■ 
■■■ 




■ 


35 














■ 






34 ■ 












• 


■ 







6.5- 
7.2 



9.3- 
9.6 



11.1- 
11.8 



12.6- 
12.9 



14.2 



14.6- 
15.3 



17.5- 
18.3 



20.8- 
21.3 



25.4- 
25.6 



Latitude 



Figure 8. Geographic variation in the number of midbody scales in Stenocercus prionotus (•) and S. caducus (■). 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., 1 /"SO' converted to 1 7.5); 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 intraspecific 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 dimoi*phic 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 pnonotus 
with the highest vertebral crests occur in 
the area of highest species density and 
documented sympatry for the ^'Ophrijoes- 
soicles 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 




















41 




















40 














■■ 






39- 






• 


• 






■■■■ 






38- 


• 




• 


• 


•• 






■ 




37- 






•• 




•••• 






1 ■ 


■ 


0) se- 
ra 

o 




•• 


• 


•• 












!! 35 

w 

a> 

^ 34 

2 


• 


• 


• 


• 


•• 
• 


■ 


■■■ 
■■■ 




■ 


® 33 

0) 


••• 
• 


•• 






••• 


• 






■■ 


31 


• 


•••• 


• 




• 










30 




•• 










■ 






29 




















28 




















27 




• 

















6.5- 
7.2 



9.3- 
9.6 



11.1- 
11.8 



12.6- 
12.9 



14.6- 
15.3 



17.5- 
18.3 



20.8- 
21.3 



25.4- 
25.6 



Latitude 



Figure 9. 
axes are 



Geographic variation in the number of vertebral crest scales in Stenocercus prionotus and S. caducus. Symbols and 
the same as Figure 8. 



contrast, in southern Peru and northern 
BoHvia the distribution of S. prionotus 
overlaps only S. scapularis and (perhaps) 
S. fimbriatus of the " Ophrijoessoides 
group," but S. prionotus is not known to 
be sympatric with either of these species.' 



' The only case of synipatry 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 "Ophryoessoicles 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 

Huanuco, Peru 
67 mm 



(b) prionotus 
Puno, Peru 
73 mm 




(c) 



prionotus 
Beni, Bolivia 
81 mm 




(d) caducus 

Sta. Cruz, Bolivia 
72 mm 




New Species of Stenocercus from Peru and Bolivia • Cadle 205 



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 (Huanuco 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- 
hriatus are known to have overlapping 
ranges; and Manu National Park in south- 
ern Peru, a reasonably accessible area 
where S. jimbriatus 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 



geographic gap between Tingo Maria and 
vicinity (Huanuco 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 without 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 associated with the se- 
ries from Explorers 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 Cashu. 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 acideatus) 
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 foUov^dng 



Figure 10. Diagrammatic representation of geographic and size-related variation in the height of the vertebral crest in Steno- 
cercus prionotus and S. caducus. Drawings were made with a camera lucida to emphasize the form and height of the vertebral 
crest. Sketches are drawn to an approximately uniform interval between the external ear opening (EE) and the white antehumeral 
stripe (AS). For each specimen the geographic location and the SVL are given (all specimens are adult males): (a) MCZ 43759, 
(b) FMNH 64799, (c) USNM 280250, (d) CM 970. Note especially the differences between the northern specimen of S. prionotus 
(a) compared to southern ones (b and c), especially given the size differences among these; and the differences between size- 
matched specimens of S. prionotus and S. caducus (b and d). 



206 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



notes on the San Juan [del Oro] locality 
from which she obtained a series of Sten- 
ocercus prionotus in the early 1950s (K. P. 
Schmidt archives, FMNH): 

Steep forest with deep undergrowth. Steep fields. 
Rainfall probably somewhat greater than at Pampa 
Grande,^ due to its colder climate and steep ex- 
posure; I have no figures. Brushy second growth 
may be burned in fairly wide patches in August, 
and [the] resort is frequently made to burning, in- 
choating a moderately dry winter period. 

As of the mid-1980s very little undisturbed 
forest was left in the vicinity of San Juan 
del Oro (personal observations). Although 
Heller provided detailed notes on some of 
the snakes and frogs from her collection, 
she makes no specific comments about the 
lizards. 

Most observations suggest that Steno- 
cercus prionotus prefers open habitats, 
such as areas of human disturbance and 
light gaps within forests (e.g., created by 
trails), rather than deep rainforests. Alter- 
natively, the observations may simply in- 
dicate the ease of observation and capture 
in more open habitats. A combination of 
cryptic coloration and escape behavior 
(rapid flight followed by immobility) pos- 
sibly makes S. prionotus very difficult to 
observe in closed-canopy rainforest, as re- 
ported for the similar species, S. fimbria- 
tus (Dixon and Soini, 1986; Avila-Pires, 
1995) and S. caducus (Scrocchi et al., 
1985).^ However, we currently lack obser- 
vations to support these statements for S. 
prionotus. 

COMPARISON OF STENOCERCUS 
PRIONOTUS WITH S. CADUCUS 

A scaly flap associated with the posthu- 
meral pockets is a unique and unquestion- 
ably derived character shared by Stenocer- 
cus prionotus and S. caducus, which sug- 
gests that these are sister species (Figs. 5, 



11).^" I am unaware of a similar structure 
in any other lizards. Some individual vari- 
ation in the precise form and size of the 
flap occurs in both species but it seems 
extremely iinprobable that these structures 
are not homologous in the two species. 

Moreoever, Stenocercus prionotus and 
S. caducus are similar in standard meristic 
characters (Table 1; Figs. 8, 9), and the 
siinilarity among Bolivian populations of 
both species has caused confusion about 
the identity of particular populations (see 
citations in the synonymy of S. prionotus) . 
Geographic, ontogenetic, or individual var- 
iation of some characters within both spe- 
cies, especially the height of the vertebral 
crest and the number of midbody scale 
rows (Fig. 8), further clouds the distinc- 
tions between them. Differentiating the 
northern populations of S. prionotus from 
S. caducus is unequivocal and facile based 
solely on the size of the vertebral crest and 
on associated meristic counts. However, 
specimens of S. prionotus from southern 
Peru and northern Bolivia are more diffi- 
cult to distinguish from S. caducus. For 
example, animals from populations of S. 
prionotus in southern Peru and Bolivia 
have less prominent vertebral crests 
(hence, higher vertebral scale counts), and 
generally lower numbers of midbody scale 
rows, than do specimens from northern 
populations (Figs. 8, 9); in these respects 
they are more similar to S. caducus. Nev- 
ertheless, even accounting for these diffi- 
culties, a combination of three qualitative 
characters is sufficient to distinguish S. 
prionotus from S. caducus, and the species 



^ I have been unable to localize Pampa Grande. 

® Cei (1993) claimed that Stenocercus caducus was 
arboreal, but Scrocchi et al. (1985) reported the be- 
havior of this species in more detail and stated that 
it was terrestrial. 



'" In my comparisons I have emphasized Bolivian 
specimens referred to Stenocercus caducus, whereas 
the type locality is "Paraguay." I have not fully con- 
vinced myself that specimens referred to this species 
from Bolivia, Paraguay, and Argentina are, in fact, all 
the same taxon. Considerable variation exists in some 
aspects of coloration and scale characters in diese 
specimens. However, my concept of S. caducus cor- 
responds to that used in current literature (e.g., Gal- 
lardo, 1959; Scrocchi et al, 1985; Cei, 1993). Only a 
thorough study of S. caducus across its range will re- 
solve this issue. 



New Species of Stenocercus from Peru and Bolivia • Codle 207 



may differ in patterns of sexual size di- 
morphism.^^ 

The Form of the Vertebral Crest. De- 
spite geographic variation in the promi- 
nence of the vertebral crest, males and fe- 
males of Stenocercus prionotiis have a dis- 
tinctly projecting serrate vertebral crest ex- 
tending from the nuchal region to the 
anterior portion of the tail (Figs. 6, 10, 12). 
The scales of the crest are strongly trian- 
gular in lateral view, are flaplike (i.e., they 
bend easily), project vertically from the 
dorsum, and are strongly differentiated 
from the adjacent dorsal scales. Although 
the crest is somewhat less developed in fe- 
males, it is prominent in both sexes. Spec- 
imens from northern Bolivia and southern 
Peru have a substantially lower crest than 
specimens from central and northern Peru 
(Fig. 10). Nonetheless, the form and pro- 
jection angle of the crest scales is the same 
as in the northern populations. 

In contrast, the scales of the vertebral 
crest in Stenocercus caducus are only mod- 
erately differentiated from adjacent dorsal 
scales in being more strongly keeled and 
mucronate. The crest in S. caducus is only 
slightly projecting in males (Fig. 10) and 
even less so in females (Fig. 12); the crest 
is mainly apparent on the neck and ante- 
rior body. In S. caducus, the scales of the 
crest are stiff and prismatic, and the main 
axis of projection is posterior rather than 
vertical, as in S. prionotiis. 

Crest height in Stenocercus varies posi- 
tively with size and thus it is critical to 
compare similar-sized specimens when 
documenting differences among popula- 
tion samples or species. This realization 
has been critical to differentiating Steno- 
cercus prionotiis from S. caducus in south- 



" I am uncertain how Fugler (1983, 1986, 1989) 
distinguished Boh\dan specimens he referred to Sten- 
ocercus aculeatus and S. caducus. In 1983 and 1986 
he referred specimens from Tumi Chucua (Beni, Bo- 
livia) to S. aculeatus. In 1989 he Usted these again, 
along with ROM specimens from San Marcos Ranch 
(Beni, Bolivia) identified as S. caducus. Fugler spec- 
imens from these localities that I have examined are 
all S. prionotus (see list of paratypes). 





Figure 1 1 . Axillary region of Stenocercus caducus showing 
the posthumeral flap (MCZ 34215). Anterior to the left. The 
posthumeral flap comprises the heavily outlined scales pos- 
teroventral to the forelimb. The opening of the posthumeral 
pocket is the heavily stippled area deep to the flap. Top, Pos- 
thumeral flap in its normal orientation covering the anterov- 
entral portion of the pocket. Bottom, the flap deflected ven- 
trally, with its posterior scales viewed from their tips. Approx- 
imately X8.5. 



ern Peru and Bolivia. Large adult males of 
S. prionotus from southern populations 
are scarce in collections. For example, al- 
though only 18 specimens of S. prionotus 
are available from northern Peru, one half 
of these are males with SVL >60 mm. In 



208 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



contrast, twice as many specimens each of 
S. prionotus and S. caducus are available 
from southern Peru and Bolivia. Yet, only 
one third of the available specimens of ei- 
ther species from these areas are males 
>60 mm SVL, and no males of S. caducus 
were >72 mm SVL. 

Figure 10 shows differences in the 
height of the vertebral crest in a series of 
males of Stenocercus prionotus from 
southern Peru and Bolivia compared with 
similar-sized specimens of S. prionotus 
from northern Peru and with S. caducus 
(the largest males of S. caducus studied 
were 72 mm SVL; see also Fig. 6). The 
trend toward lower crests in S. prionotus 
from the southern part of its range is evi- 
dent, as is the difference between S. cad- 
ucus and S. prionotus. A comparison of all 
specimens suggests that the difference in 
the height of the vertebral crest between 
S. prionotus and S. caducus males begins 
to be apparent by approximately 65 mm 
SVL and becomes pronounced at around 
70-75 mm SVL. No males of S. caducus 
>72 mm SVL were among the specimens 
examined, despite the availability of a large 
number of specimens froin southern Bo- 
livia, including a series of 31 specimens 
(10 males >60 inm SVL) from the vicinity 
of Santa Cruz. A siniilar contrast in crest 
height appears in females of the two spe- 
cies (Fig. 12). 

Angulate Temporal Scales. Stenocercus 
prionotus has two (occasionally three) very 
strongly keeled, projecting angulate tein- 
poral scales on each side (Fig. 4). These 
are much larger than adjacent scales on 
the head and they are partially or com- 
pletely separated from the large posterior 
head scales (parietals, postparietals, and 
occipital) by one row of small keeled scales 
(occasionally partially doubled). Stenocer- 
cus caducus usually has two (occasionally 
three) angulate temporals that are smaller 
and less projecting than those in S. prion- 
otus. In S. caducus the angulate temporals 
may or may not be larger than adjacent 
posterior head scales and they are not 




Figure 12. Size-matched females of Stenocercus prionotus 
and S. caducus from Bolivia. Top, S. prionotus (USNM 
269022, snout-vent length [SVL] 91 mm). Bottom, S. caducus 
(UTA 38046, SVL 93 mm). Note the subtle difference in crest 
height between the two specimens and their otherwise similar 
patterns. 



strongly differentiated from other posteri- 
or head scales. 

Color Pattern of the Gular Region. 
Many specimens of Stenocercus prionotus 
have a regular pattern of alternating diag- 
onal light and dark stripes on the throat. 
These usually converge closely toward the 
midline (Fig. 7) and are most easily visu- 
alized in preseived specimens submerged 
in alcohol. This pattern consists of a dark 
stripe beginning at a point on the lower 
labials in line with, but broader than, the 
subocular dark bar. The stripe projects 
posteromedially, gradually fading and 
blending with the ventral ground color on 
the neck anterior to the pectoral region. 
The dark stripe is bordered on either side 
by a distinct pale stripe. Anteriorly, this se- 
ries is preceded by another dark and an- 
other pale stripe. The dark stripes are usu- 
ally approximately twice as wide as the 
pale ones, although not always (e.g., the 
dark stripes are only slightly wider than 
the pale ones in MCZ 150243). In life the 
pattern may manifest itself as a series of 
pale stripes on a darker background (e.g., 
the "gular area streaked by several light 



New Species of Stenocercus from Peru and Bolivia • Cadle 



209 



cream colored lines" in the life colors of 
the holotype). 

The gular region appears uniform in 
many preserved specimens of Stenocercus 
prionotus, but I suspect this is a preser- 
vation artifact. Occasional specimens have 
pale spots in the pectoral region, and oth- 
ers are essentially unicolor and without ap- 
parent pattern (again, probably a preser- 
vation artifact). 

On the other hand, the throat pattern of 
Stenocercus caducus is highly variable and 
irregular. When a distinctive pattern is 
present, it most often consists of light 
spots rather than alternating stripes (Fig. 
13). Cope (1862) described the holotype 
of S. caducus from Paraguay as having a 
dark throat that was "light varied" (i.e., 
variegated, or spotted), and some speci- 
mens I examined have this pattern (Fig. 
13). None of several color descriptions for 
Argentinian specimens of S. caducus men- 
tion stripes or spots on the throat. Scrocchi 
et al. (1985) described living examples as 
having pale spots in parallel transverse 
rows in the pectoral region or with pale 
spots on the abdomen, but did not com- 
ment on the throat pattern; Gallardo 
(1959) described the ventral coloration as 
"pale olive with some scattered pale spots; 
throat darker"; and Cei (1993) described 
the venter as "dark brownish with series of 
rounded pale spots, sometimes anastomos- 
ing along the length of a median line." Al- 
though no authors mention alternating 
light and dark stripes on the throat in S. 
caducus, UTA 38046 does have this pat- 
tern (Fig. 13). But in this specimen the 
stripes are confined to the lateral edges of 
the throat (i.e., do not closely approach the 
midline as in S. prionotus). Apart from the 
throat pattern, the coloration of S. prion- 
otus and S. caducus seems to be veiy sim- 
ilar judging from descriptions of S. cadu- 
cus in the literature (Gallardo, 1959; 
Scrocchi et al, 1985; Cei, 1993). 

A Possible Dijference in Sexual Size Di- 
morphism. Data presented in Table 1 sug- 
gests another contrast between Stenocer- 
cus prionotus and S. caducus: S. prionotus 




Figure 13. Gular patterns in Stenocercus caducus. Top, typ- 
ical throat pattern consisting of light spots on a dark back- 
ground (BMNH 1927.8.1.163). Bottom, variant pattern consist- 
ing of stripes confined to the lateral portion of the throat (UTA 
38046). Connpare to Figure 7. 



210 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



is not dimorphic in the maximum sizes at- 
tained by males and females, whereas fe- 
males of S. caducus apparently attain 
about 20 mm greater SVL than males. 
However, estimating maximum size is sub- 
ject to considerable sampling error so this 
distinction should be considered to be only 
provisional. Nonetheless, males of S. 
prionotus with SVL >80 mm are known 
from the northern (USNM 193685) and 
southern (USNM 280250, BMNH 
98.6.9.4) portions of its range, even though 
the three largest males from the largest 
population sample (23 specimens in 
FMNH from Puno Department, Peru) 
had an SVL of 73 mm (this sample includ- 
ed six adult females with an SVL of 78-89 
mm). 

In contrast, a sample of 39 Stenocercus 
caducus from Bolivia included 12 adult 
males, none of which had an SVL >72 
mm; in the total sample of S. caducus iJSJ 
= 43) 17 females had an SVL >80 mm 
(range 80-93 mm).^- Thus, unless a sys- 
tematic collecting bias against males exists, 
the different pattern of sexual size dimor- 
phism provides another character distin- 
guishing S. prionotus and S. caducus. Data 
presented in Table 1 suggests that other 
species of the " Ophrijoessoides group" 
may be size dimoi*phic (S. fimbriatus, S. 
hiiancabambae, and ?S. scapularis) or not 
(S. aculeatus), and either males (S. huan- 
cahambae) or females (S. caducus and S. 
fimbriatus) may attain a larger body size. 

DISTRIBUTIONS OF STENOCERCUS 
PRIONOTUS AND S. CADUCUS IN 
EASTERN BOLIVIA 

The ranges of Stenocercus prionotus 
and S. caducus approach one another in 



eastern Bolivia, but do not overlap. ^^ Cur- 
rently, the two closest documented locali- 
ties are, for S. prionotus, near the junction 
of the Rio Madidi and the Rio Beni 
(BMNH 98.6.9.4), and for S. caducus, 
many specimens from the vicinity of Santa 
Cruz de la Sierra (see above discussion for 
S. prionotus and Appendix; Fig. 1). Sten- 
ocercus caducus is also known from the 
Bolivia— Brazil border in the region of the 
Serrania de Huanchaca in northern Santa 
Cruz Department, Bolivia. ^^ 

I am aware of no specimens of Steno- 
cercus referable to either S. caducus or S. 
prionotus between the Rio Beni valley and 
roughly a line connecting Santa Cnaz and 
the vicinity of Serrania de Huanchaca. 
Southeast of the Rio Beni, the central part 
of the Beni basin (the Llanos de Mojos) is 
characterized by flooded savanna grass- 
lands, palm savannas, swamps, and other 
habitats that are inundated for significant 
portions of the year; terra firme forests are 
restricted to somew^hat elevated levees 
along rivers (Clapperton, 1993: 196; Han- 
agarth, 1993). Stenocercus prionotus or S. 
caducus seem unlikely to occur in this area 
except possibly in these galleiy forests, al- 
though Fugler (1986) reported S. priono- 
tus (as Ophryoessoides aculeatus) in sea- 
sonally flooded forest during the dry sea- 
son. 

Stenocercus prionotus and S. caducus 
probably are segregated by habitat in Bo- 
livia and their distributions may not over- 
lap. Stenocercus prionotus is associated 



^- These sizes are somewhat larger than those pre- 
viously reported (81 mm; Gallardo, 1959; Scrocchi et 
al, 1985; Marcus, 1986). Cei (1993) stated that S. 
caducus reaches only 75 mm SVL in Argentina. Sexes 
were not given for any individual or sexed specimens 
in these reports. 



'"^ All references to "Ophryoessoides aculeatus" in 
Bolivia (e.g., Fugler, 1983, 1986) that I verified have 
referred to Stenocercus prionotus. However, given 
the general confusion of species in this complex, 
some records not traced will have to be checked to 
rule out the possibility that they do not refer to S. 
caducus or perhaps some other species of the 
"Ophryoessoides group," such as S. fimbriatus or S. 
scapidaris (see Distribution Patterns in Stenocercus 
prionotus). 

i-i See the Appendix, UTA 38048. Michael Hawey 
(personal communication) recently obtained speci- 
mens of Stenocercus caducus at El Refugio, a lowland 
locality at the southern end of the Serrania de Huan- 
chaca (14°44'S. 61°01'W). 



New Species of Stenocercus from Peru and Bolivia • Cadle 



211 



with upper Amazonian and lower montane 
rainforests with annual rainfall greater 
than 2,000 mm in both Peru and Bolivia. 
On the other hand, confirmed localities of 
S. caducus are within the physiographic 
domain broadly referred to as chaco, in- 
cluding a mixture of diy forests, palm sa- 
vannas, galleiy forests, deciduous forests, 
and ecotonal areas (Scrocchi et al., 1985; 
Marcus, 1986; Cei, 1993). Short (1975) 
and Parker et al. (1993) described the di- 
versity of chaco habitats. Gallardo (1979: 
table 12.1) listed S. caducus as a species 
"basically restricted to the chaco." Average 
annual rainfall in this area is less than 
1,000 mm. Stenocercus caclucus is known 
from Parque Nacional Noel Kempff Mer- 
cado and vicinity in Bolivia (see footnote 
14; Haivey, 1998). This area is character- 
ized by a complex mixture of habitat types, 
including deciduous forests and cerrado 
enclaves, and with an annual rainfall of 
1,400-1,500 mm (Killeen, 1998). Hai-vey 
(1998) encountered S. caducus at granitic 
outcrops covered by semideciduous forests 
and more open habitats. The herpetofauna 
of this site is a mixture of species that are 
typical of Amazonian and of chaco envi- 
ronments (Hai^vey, 1998; personal obser- 
vations). 

The range of Stenocercus caducus ex- 
tends outside the strictly defined chaco re- 
gion (see Short, 1975, and Clapperton 
1993, for discussion) on the southeastern 
edge of its range east of the Rio Paraguay 
and in the Andean foothills of southern 
Bolivia and northern Argentina (Fig. 1). 
Harvey (1997) reported S. caducus from 
"subtropical wet forests" (1,150—2,050 m 
elevation) in southern Bolivia. He charac- 
terized S. caducus as a "Chacoan species 
that invade[s] the Andean foothills . . . [in- 
cluding] those distributed within the Gran 
Chaco or that occur in diy forests sur- 
rounding the Gran Chaco" (Hai-vey, 1997: 
35). The montane wet forests (yungas) of 
this area are restricted to ridges high 
enough for cloud formation during much 
of the year (generally > 1,500 m elevation), 
and they are surrounded by deciduous dry 



forest (Schulenberg et al., 1997). The cli- 
mate of this area is generally dry and it 
receives only about 1,200 mm of rainfall 
per year (Hoist, 1997). 

The transition between the wet rainfo- 
rests of Peru and northern Bolivia (range 
of Stenocercus prionotus) and the chaco 
habitats (range of S. caducus) occurs in a 
very broad ecotone consisting of savannas, 
evergreen shrublands, and gallery forests 
of the Beni basin and Rio Mamore drain- 
age, from which no specimens of either S. 
prionotus or S. caducus have been report- 
ed. The piedmont forests of the Andes be- 
tween the known ranges of S. prionotus 
and S. caducus, which are wetter than ad- 
jacent lowland forests because of the mod- 
erating effect of the Andes, provide one 
potential route for contact or overlap of 
their ranges. 

The eastern distributional limits of Sten- 
ocercus caducus along the Bolivia— Brazil 
frontier are not well understood. I am un- 
aware of verified records from Brazil, al- 
though the species does occur close to the 
Brazilian border in the vicinity of the Ser- 
rania de Huanchaca in Parque Nacional 
Noel Kempff Mercado. Some references 
to "Stenocercus caducus" from western 
Brazil (e.g., Mato Grosso State; Cope, 
1887; Boulenger, 1903) likely refer instead 
to an undescribed species veiy similar to 
S. caducus (P. E. Vanzolini and E. E. Wil- 
liams, personal communication; personal 
observations). However, the ranges of S. 
caducus and the undescribed species in 
eastern Bolivia-Paraguay and western Bra- 
zil are not well defined; the two species 
may be separated by the seasonally inun- 
dated savannas of the pantanal. Addition- 
ally, few specimens of S. caducus appar- 
ently exist from the chaco of northwestern 
Paraguay, although Aquino et al. (1996) re- 
ported specimens from Parque Nacional 
Defensores del Chaco (approximately 
20°30'S, 60°20'W), as well as other Para- 
guayan localities in more mesic regions 
east of the Rio Paraguay. 



212 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



IS THE DISTRIBUTION OF 
STENOCERCUS ACULEATUS 
DISJUNCT? 

In the process of diagnosing Stenocercus 
prionotus I reviewed the characters and 
distribution of S. aculeatus. In addition to 
variation in some characters of uncertain 
significance, some aspects of the distribu- 
tion of S. aculeatus seem pecuHar (Fig. 2). 
First, the distribution of S. aculeatus ap- 
pears to be disjunct between northern 
Peru and eastern Ecuador. Although the 
type locality is in northern Peru (Moya- 
bamba, San Martin Department), most 
specimens are from eastern Ecuador (Fig. 
2). The two areas from which specimens 
are known (Fig. 2) are separated by a 
broad geographic gap through which 
courses the main tributary of the upper 
Amazon, the Rio Marafion. Neither S. acu- 
leatus nor any other species of Stenocercus 
was obtained during hei"petofaunal surveys 
in northern Loreto Department, Peru 
(Duellman and Mendelson, 1995), north- 
ern Amazonas Department, Peru (Rio Ce- 
nepa and Rio Santiago; J. E. Cadle and R. 
W. McDiarmid, unpublished data), or dur- 
ing a rapid biological assessment of the 
Cordillera del Condor region of southeast- 
ern Ecuador and northern Peru (Schulen- 
berg and Awbrey, 1997a). Stenocercus acu- 
leatus is known from many localities in ad- 
jacent regions of Ecuador. 

Second, all Ecuadorian localities for 
Stenocercus aculeatus are in the drainages 
of the Rio Pastaza and the Rio Curaray. 
No specimens are known from the Rio 
Napo drainage just to the north, even 
though no recognized physiographic or 
faunal break seems to separate the Rio 
Napo drainage from the Rio Curaray 
drainage. However, all localities that have 
been sampled comprehensively from the 
Rio Napo are on the left (northern) bank 
of the river (e.g., Duellman, 1978; Vitt and 
De la Torre, 1996; unpublished list from a 
large collection from the Jatun Sacha Bi- 
ological Station assembled and under 
study by Gregoiy Vigle). The absence of 



S. aculeatus from Santa Cecilia (Duell- 
man, 1978) is probably real rather than 
sampling error, given the intensity of col- 
lecting over several years at the site. Thus, 
S. aculeatus possibly does occur on the 
right (south) bank of the Rio Napo and 
will be recorded once large collections are 
made there. 

The apparent geographic disjunction of 
Stenocercus aculeatus between northern 
Peru and eastern Ecuador may correspond 
to some character differences among sam- 
ples that should be studied more thor- 
oughly (Cadle, unpublished data). For ex- 
ample, Peruvian specimens of S. aculeatus 
have veiy deep postfemoral pockets (Type 
5) in both sexes, whereas the postfemoral 
pockets are more weakly developed in 
specimens from Ecuador (Type 2 or 3 in 
both sexes). Ecuadorian specimens also 
appear to have more scales in the vertebral 
row and fewer subdigital scales on the 
fourth toe than do Peruvian specimens. All 
of these impressions are based on small 
sample sizes (Appendix). 

The significance of diese differences is 
unclear without a more detailed study of 
variation among populations of Stenocer- 
cus aculeatus. However, one possibility is 
that two or more species are represented 
in specimens currently referred to S. acu- 
leatus, in which case the distributions of 
individual taxa may be not be contiguous. 
This is analogous to the previous confusion 
of S. finihriatus and S. prionotus with S. 
aculeatus. Taxonomic recognition of S. 
finihriatus and S. prionotus has concomi- 
tantly reduced the geographic distribution 
understood for S. aculeatus. Consequently, 
a more comprehensive systematic analysis 
of S. aculeatus with special reference to a 
comparison of Ecuadorian and Peruvian 
populations is warranted. If two species 
are recognized, the name Liocephalus an- 
gulifer Werner (1901) is available for the^ 
Ecuadorian populations. 

Key to Species of the "Ophryoessoides 
group" of Stenocercus 

Because of the general confusion about 
the species considered herein (e.g., see the 



New Species of Stenocercus from Peru and Bolivia • Codle 213 



synonymy of Stenocercus prionohis) , I pro- 
vide the following key as a guide for iden- 
tifications. The key will work for those spe- 
cies of Stenocercus in Peru or Bolivia with 
keeled ventral scales, enlarged posterior 
head scales, and one row of moderately to 
greatly enlarged supraoculars COphryoes- 
soides group" as used herein). I have also 
included the three other currently recog- 
nized species having these characteristics, 
S. enjthrogaster (Hallowell), S. dumerilii 
(Steindachner), and S. tricristatus (Du- 
meril), although these are not known from 
Peru or Bolivia and are unlikely to occur 
there. Character and distributional data in 
the key for S. dunierilii and S. tricristatus 
follow Avila-Pires (1995). 

I also include in the key an undescribed 
species with keeled ventrals and enlarged 
head plates and supraoculars from Ama- 
zonas Department, Peru, but I am un- 
aware of other undescribed species of the 
"Ophrijoessoides group" from Peru or Bo- 
livia. However, an undescribed species 
similar to Stenocercus caducus (but lacking 
a posthumeral flap) is known from western 
Brazil (Mato Grosso) and is not included 
in the key. Additional study of S. iridescens 
from the Pacfic lowlands of Peru and Ec- 
uador is needed (Cadle, 1998: footnote 4) 
and, as indicated above, a thorough mod- 
ern study of variation in S. aculeatus (Am- 
azonian Ecuador and Peru) is also war- 
ranted. Other undescribed species may re- 
side within either of these named taxa. 
The key Mdll permit identification of all Pe- 
ruvian and Bolivian taxa previously con- 
fused with S. aculeatus (e.g., DLxon and 
Soini, 1975, 1986 [S. Jimbriatus]; Fugler, 
1983, 1986, 1989 [S. prionotus]) and S. ir- 
idescens (e.g., S. huancahambae and S. 
limitaris; see Cadle, 1991, 1998). The key 
also should work for Ecuadorian species, 
with the caveat that I have paid less atten- 
tion to Ecuadorian Stenocercus except as 
necessary in conjunction with work on Pe- 
ruvian species. Of the species covered, 
only S. acideatus, S. iridescens, and S. lim- 
itaris are definitely known from Ecuador. 



The key should be viewed as a means of 
identifying a set of phenotypically similar, 
but not necessarily closely related, species 
within Stenocercus in the broad sense. All 
other species of Stenocercus in Peru and 
Bolivia have smooth (or at most only very 
weakly keeled) ventrals and more frag- 
mented supraoculars and head plates; see 
Fritts (1974), Frost (1992), and Cadle 
(1991, 1998) for discussion and illustra- 
tions. Many of these species also have 
granular scales on the body or posterior 
surface of the thigh, neither of which is 
present in species covered by the key. The 
keys and discussions in Fritts (1974) and 
Cadle (1991, 1998) are useful for identi- 
fying these other species. 

The key assumes familiarity with char- 
acters of the mite pockets, head scales, and 
neck folds and crests outlined in Cadle 
(1991) (see also Materials and Methods). 
In most cases I have used characters that 
show minimal sexual dimoqDhism so that 
specimens of either sex can be identified; 
exceptions are noted. It is useful to keep 
in mind that, in most species of Stenocer- 
cus, scales of juveniles are more promi- 
nently keeled than in adults, even when 
the corresponding scales of adults, such as 
head scales and dorsal body scales, are 
smooth. Instances of possible confusion in 
the key are indicated. The extent of de- 
velopment of posthumeral and postfemor- 
al mite pockets varies according to sex and 
size in many species of Stenocercus, al- 
though such variation seems less extensive 
in this set of species than in many others; 
I have indicated the range of variation in- 
cluding juveniles and adults of both sexes 
in the key. Summaiy geographic distribu- 
tions are given for each species as a rough 
guide to known occurrences. However, 
these should be used cautiously as ancil- 
laiy information in identifying specimens 
because distributions of species are some- 
times poorly circumscribed. For greatest 
utilit}^ the key should be used in conjunc- 
tion with illustrations herein and in Cadle 
(1991, 1998) and Avila-Pires (1995). 



214 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



1. Canthal and supraciliaries forming a pro- 

nounced crest that ends in an enlarged, 
erect, postsupraciliaiy that may be dis- 
tinctly pointed or blunt. Posthumeral and 
postfemoral mite pockets absent (Type 1 

in both instances) 2 

Cantlials and supraciliaries not forming a 
pronounced crest; no enlarged, erect post- 
supraciliaiy. Posthumeral pocket absent 
(Type 1) to deep (Type 4). Postfemoral 
pocket absent (Type 1) to deep (Type 5) 
3 

2. Enlarged postsupraciliary distinctly pointed. 

Two enlarged scales above ear opening. 
Tibia approximately equal to thigh length 

Stenocercus dumerilii (Steindachner) 

(northeastern Para, Brazil) 
Enlarged postsupraciliary blunt. No enlarged 
scales above ear opening; tibia distinctly 

shorter than thigh 

Stenocercus tricristatus (Dumeril) 

(known only from the holotype, probably 
from the state of Minas Gerais, Brazil) 

3. Superciliary scales projecting laterally shelf- 

like above the orbit in adults, rectangular 
in dorsal view.'^' Midbody dorsal scale rows 
more than 55 (59-70). Postfemoral pocket 

well developed (Type 3 or 5) 

Stenocercus scapularis (Boulenger) 

(intermediate elevations on the Andean 
slopes of central and southern Peru; known 
elevations greater than 1,000 m) 
Superciliaiy scales not projecting laterally, 
the anterior ones elongate, strongly over- 
lapping. Midbody dorsal scale rows fewer 
than 55 (30—53). Postfemoral pocket vari- 
able (Type 1, 2, 3, or 5) 4 

4. Posthumeral mite pocket deep (TjqDe 4) and 

with an associated scaly flap extending 
from its anteroventral border and partally 

concealing it 5 

Posthumeral mite pocket absent to deep 
(Type 1, 2, 3, or 4) but without an asso- 
ciated flap 6 

5. Vertebral crest strongly projecting in both 

sexes, serrate, extending from the nuchal 
region to the proximal portion of the tail; 
its individual scales triangular, flaplike. 
Throat often with alternating oblique dark 
and light stripes. Two enlarged, strongly 
keeled and projecting angulate temporal 



'' The superciliaries in juveniles of Stenocercus sca- 
pularis have a more typical shape. The extent of 
shelf-like projection and change to a more rectan- 
gular shape seem positively correlated with body size 
and thus develop with age. The number of dorsal 
scale rows and the development of the postfemoral 
pocket are useful clues for subadults. 



scales on each side 

Stenocercus prionotiis Cadle 

(rainforested lowlands and Andean foothills 
of eastern Peru and northern Bolivia) 
Vertebral crest low, nonprojecting in both 
sexes (slightly higher in males), evident 
mainly on neck and anterior body; its in- 
dividual scales prismatic and lying more or 
less flat. Throat pattern variable, but usu- 
ally consisting of light spots on a darker 
background when evident. Angulate tem- 
poral scales keeled, but not greatly en- 
larged, and nonprojecting 

Stenocercus caducus (Cope) 

(deciduous woodlands and ecotonal areas of 

southern Bolivia, northern Argentina, and 

the chaco of Paraguay) 

6. A fringe of enlarged fimbriate scales on the 

distal posterodorsal surface of thigh. Sev- 
eral longitudinally oblique rows of large, 
strongly keeled scales on shank.''' Scales 
between vertebral and dorsolateral crests 
smooth or occasionally veiy weakly keeled. 

.-. Stenocercus finibriatus Avila-Pires 

(lowlands of eastern Peru and central western 
Amazonian Brazil) 
No fringe of fimbriate scales on thigh or 
strongly keeled oblique scales on shank. 
Scales between vertebral and dorsolateral 
crests moderately to strongly keeled at 
least posteriorly; dorsolateral crest may be 
weakly developed, but dorsal scales still 
strongly keeled 7 

7. Posthumeral mite pocket variable (Type 1, 2, 

3, or 4). Postfemoral mite pocket variable 
(Type 1, 2, 3, or 5). Head scales smooth 
or keeled. Angulate temporal scales 
keeled, may be projecting and bladelike. 
Internasals usually 4 or more (occasionally 
3, never 2), often irregular in pattern and 

shape. 8 

Posthumeral mite pocket absent or weakly 
developed (Type 1 or 2). Postfemoral mite 
pocket absent (Type 1). Head scales 
smooth. Angulate temporal scales smooth; 
none bladelike and projecting. Two polyg- 
onal internasals in contact on the midline, 
each broader laterally than medially. 

Stenocercus iridescens (Giinther) 

(Pacific lowlands and intermediate elevations 
of western Ecuador and northwestern 

Peru) 

8. One to 3 strongly keeled, but nonprojecting, 

angulate temporal scales in line with the 
superciliary row between the lateral tem- 



"^ The fimbriate scales form a projecting fringe on 
the distal portion of the thigh. Both the fimbriate 
scales and the oblique scales on the shank are rela- 
tively more prominent in juveniles than adults. 



New Species of Stenocercus from Peru and Bolivia • Cadle 215 



porals and the posterior dorsal head scales. 
Two subequal can thai scales on each side. 

Head scales keeled, at least posteriorly 9 

Two projecting bladelike angulate temporals 
in line with superciliary row. A single can- 
thai on each side (rarely, 2 are present but 
in that case 1 is much larger than the oth- 
er). Head scales smooth or keeled. 11 

9. Posthumeral pocket moderately developed 

in males (Type 2 or 3), absent in females 
(Type 1). Postfemoral pocket absent in fe- 
males (Type 1), moderate to deep in males 
(Type 3 or 5). Anterior gular scales weakly 

to strongly keeled. 

Stenocercus erythrogaster (Hallowell) 

(northern Colombia) 
Posthumeral and postfemoral pockets deep 
in both sexes (Types 4 and 5, respective- 
ly).'' Anterior gular scales smooth to weak- 
ly keeled. 10 

10. Inteiparietal indistinct, parietal eye not visi- 

ble. Three occipitals. Dark subocular bar 
absent. Three angulate temporals separat- 
ed from large posterior head scales by a 

row of tiny scales Stenocercus new species 

(known from a single specimen [Appendix] 
from the inter-Andean valley of the Rio 
Maranon near Balsas, Amazonas Depart- 
ment, Peru) 
Inteiparietal distinct, parietal eye visible. 
Two occipitals. Dark subocular bar pre- 
sent. One angulate temporal much larger 
than others and in contact with at least 1 

other enlarged posterior head scale. 

Stenocercus liniitaris Cadle 

(intermediate elevations [600—2,200 m] of the 
Andes on the Pacific versant of southwest- 
ern Ecuador and northwestern Peru) 

11. Head scales smooth to slightly wrinkled in 

adults; weakly keeled, wrinkled, or rugose 
in juveniles. Prominent dorsolateral crest 
on body from neck to base of tail and con- 
tinuous with both supra-auricular crest 
and antehumeral crest. Postfemoral pock- 
et moderate to deep (Type 2, 3, or 5). 

Stenocercus aculeatus (O'Shaughnessy) 

(rainforested lowlands and intermediate ele- 
vations of northern Peru adjacent to the 
Andes and in eastern Ecuador) 
Head scales strongly keeled or multicarinate 
in juveniles and adults. Dorsolateral crest, 
when present, weak and restricted to neck 
and anterior body. Postfemoral pocket 

deep (Type 5). 

Stenocercus huancahanibae Cadle 



'' An undescribed species in the ne.xt couplet of the 
key is known only from a single adult male. The dis- 
tributions of species in couplets 9 and 10 should be 
used as ancillaiy data for identification. 



(diy inter-Andean valleys of the upper Rio 
Maraiion in Cajamarca and west central 
Amazonas departments, northern Peru) 

ACKNOWLEDGMENTS 

Loans and other assistance were facili- 
tated by Linda Ford, Barrel Frost, and 
Charles W. Myers (AMNH); E. Nicholas 
Arnold and Colin J. McCarthy (BMNH); 
John Wiens (CM); Cassy Redhed, Alan 
Resetar, and Harold Voris (FMNH); Wil- 
liam E. Duellman, Christopher J. Raxwor- 
thy, and John E. Simmons (KU); Frank 
Burbrink and Douglas Rossman 
(LSUMNS); Ross MacCulloch and Robert 
W. Murphy (ROM); Roy W. McDiarmid, 
Steven W. Gotte, W. Ronald Heyer, and 
Robert R Reynolds (USNM); and Jona- 
than Campbell and Michael B. Harvey 
(UTA). Victor Morales permitted me to ex- 
amine a specimen of Stenocercus fimbria- 
tus in his care. I am grateful to Wade C. 
Sherbrooke for providing copies of his 
field notes and other information on spec- 
imens he collected. I owe a great debt to 
the late Ernest E. Williams, who was ex- 
tremely generous with discussion, notes, il- 
lustrations, and encouragement. Williams, 
Paulo E. Vanzolini, and Richard Etheridge 
long ago distinguished Stenocercus prion- 
otus and two other species I described (S. 
huancabainbae and S. liniitaris) but kindly 
let my work on the group unfold with their 
gracious consent. Vanzolini supplied a 
copy of a portion of Balzan (1931), helped 
interpret Balzan's localities, and pointed 
out Metraux's work to me. I am indebted 
to several people for the special efforts 
they made in tracking down information 
about particular collections: Bruce Patter- 
son (FMNH) supplied information on the 
collections of Colin Sanborn and Hilda 
Heller from Puno Department, Peru; Alan 
Resetar and Cassy Redhed (FMNH) dug 
into the Schmidt archives and found ad- 
ditional information about Hellers collec- 
tion; Charles W. Myers (AMNH) did the 
same for Haivey Basslers journeys in 
northern Peru and provided the base map 
used to prepare Figure 1; Robert S. Voss 



216 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



checked AMNH sources for information 
on Keays's Peruvian localities; and Colin J. 
McCarthy (BMNH) clarified the confusion 
about P. O. Simons's "Palca" locality, if not 
the locality itself. Robin Andrews, Tom 
Jenssen, and A. Stanley Rand discussed as- 
pects of geographic variation and the uses 
of vertebral crests in S. prionotus with me. 
Laszlo Meszoly drew Figures 4, 5, and 11. 
For comments on the manuscript I thank 
Richard Etheridge and Charles W. Myers. 
The research was supported in part by a 
faculty grant from the School of Arts and 
Sciences of Harvard University; publica- 
tion costs were supported by the Colles 
Fund of the MCZ. 

APPENDIX: SPECIMENS EXAMINED 

Institutional abbreviations are as fol- 
lows: 

AMNH American Museum of Natural 

History, New York 

ANSP Academy of Natural Sciences 

of Philadelphia 

BMNH The Natural History Muse- 

um, London 

CM Carnegie Museum of Natural 

History, Pittsburgh, Pennsyl- 
vania 

FMNH The Field Museum, Chicago 

KU Natural History Museum, 

University of Kansas, Law- 
rence 

LSUMNS Louisiana State University 
Museum of Natural Science, 
Baton Rouge 

MCZ Museum of Comparative Zo- 

ology, Harvard University, 
Cambridge 

ROM Royal Ontario Museum, To- 

ronto 

USNM National Museum of Natural 

History, Washington, D.C. 

UTA University of Texas at Arling- 

ton 

Bracketed information was inferred 
from sources listed in the Materials and 
Methods. For specimens of other species 
of the "Ophryoessoides group" examined 



(S. huancabamhae, S. iridescens, and S. 
limitaris), see Cadle (1991, 1998). Bold- 
face numbered localities 1—4 are known or 
suspected areas of sympatry between the 
species listed and Stenocercus prionotus. 
They correspond to numbered localities in 
Figures 1 and 2 and in the text discussion. 

Stenocercus aculeatus 

ECUADOR: Prov. Morona-Santiago: Chiguaza 

[ca. 1,000 m; 01°59'S, 77°58'W] (USNM 200882-84). 
[?Prov. Napo/Tungurahua]: Llanganates area'** 
(FMNH 2.3527). Prov. Pastaza: Rio Pastaza, Abita- 
gua [01°23'S, 78°05'W] (FMNH 2.5803-05, 26892, 
28011, 28057 [ = 17 specimens]). Rio Pastaza, Alpay- 
acu [01°28'S, 78°07'W] (FMNH 3926-27; MCZ 
8081). Canelos [01°35'S, 77°45'W; 530 m] (MCZ 
38530). Montalvo, Rio Robonaza [314 m; 02°04'S, 
76°58'W] (USNM 200892). [?Prov. Pastaza]: Ranos, 
Mera Trail [? = between Raiios and Mera'^; approx- 
imately 01°30'S, 78°10'W] (FMNH 28012). 

PERU: [Depto. La Libertad]: E Peru, Pampa 
Seca, Rio MLxiolla [ = Rio Mishollo] Valley, Upper 
Huallaga region, 4300 ft [2, 1,311 m; approximately 
08°16'S, 76°58'W]2« (AMNH 57085). [Depto. Lor- 



'' I have been unable to localize this. The Cordil- 
lera de los Llanganates is a high range (to >4,500 m) 
in the Cordillera Oriental north of the Rio Pastaza 
(Paynter, 199.3). The locality may refer to lower ele- 
vations in this range. 

"^ Rafios is a famous collecting locality at the foot 
of Volcan Tunguraliua at 1,820 m. That would be an 
altitudinal record for Stenocercus aculeatus. I inter- 
pret the locality as stated in the FMNH catalogues 
as being on the trail between Raiios and Mera, which 
is at 1,160 m. See Rrown (1941) and Chapman (1926) 
for discussion. 

-" Hai"vey Rassler collected Stenocercus for the 
AMNH at two localities on the Rio Mixiolla ( = Rio 
Mishollo): Pampa Seca and La Pinita (see Stenocer- 
cus finibriatus), as listed in AMNH catalogues. The 
Rio Mishollo originates in southeastern La Libertad 
Department, flows eastward, and joins the Rio Hual- 
laga in southwestern San Martin Department. The 
elevations given for these localities, 1,067 m and 
1,311 m, indicate that they lie in the narrow stretch 
of the valley that straddles the boundary between La 
Libertad and San Martin departments (departmental 
maps produced by the Instituto Geografico Nacional, 
Lima). I identify these localities as Pampaseca and 
Piiiita, respectively, in extreme southeastern La Lib- 
ertad Department, as indexed by Stiglich (1922). 
Roth localities are in Ongon District and the coor- 
dinates given are those for the town of Ongon. Stig- 
lich (1922) states that Piiiita is a small village on the 
Quebrada Pedernal, a left tributary of the Rfo Mish- 
ollo. Apart from indicating that Pampaseca is a farm. 



New Species of Stenocercus from Peru and Bolivia • Cadle 217 



ETo]: NE Peru, Front Range between Moyabamba 
and Cahuapanas. 3000 ft [915 m; approximately 
05°37'S, 77°00'W] (AMNH 57083). Northeastern 
Peru: Icuta on Balsapuerto-Moyabamba trail, 3500 ft 
[1,067 m; 05°58'S, 76°40'W; given as "Icuto" or "len- 
to Cuesta" by Lamas, 1976] (AMNH 56413). 

Stenocercus caducus 

BOLIVIA: No specific locality: BMNH 

1946.8.29.76 (holotype of Leiocephahis bolivianus 
Boulenger); CM 4583-84. Depto. Chuquisaca: Sud 
Cinti, trail from Rinconada Bufete to El Palmar 
[1,170-2,000 m; approximately 20°50'S, 64°21'W] 
(UTA 39102). Depto. Santa Cruz: Buena Vista, ca. 
500 m [17°27'S, 63°40'W] (MCZ 20625-26, 29023; 
FMNH 16165, 21486, 21511; BMNH 1927.8.1.163- 
164; CM 4527, 4550-51, 4558, 4587-88, 4605, 4607, 
4616, 4626, 4634-36, 4641). Las Yuntas [ = Las Jun- 
tas;-' 18°38'S, 63°08'W] (CM 970). Provincia Chiqui- 
tos, Santiago (Serrania and nearby), 700-750 m 
[18°19'S, 59°34'W] (FMNH 195983). [Provincial 
Chiquitos, Canton El Cerro, Finca Dos Milanos, 
17°27'30"S, 62°20'00"W (UTA 38046). Provincia Sara, 
eastern Bolivia, 600 m [17°27'S, 63°40'W] (BMNH 
1907.10.31.7-8). Provincia Sara, Santa Cruz de la Si- 
erra [17°48'S, 63°10'W] (CM 966, 969, 13018). Prov- 
incia Sara, Rio Surutu W of Buena Vista [17°24'S, 
63°51'W] (CM 4590). Provincia Sara, Rio Colorado 
[17°38'S, 63°54'W] (CM 4598). [Provincia] Velasco, 
Inselbergs near Florida [14°38'S, 6ri5'W] (UTA 
38048). [Depto. Takija]: Villa Monies [21°15'S, 
63°30'W] (MCZ 28634). Misidn San Francisco 
[21°15'S, 63°30'W] (BMNH 98.7.7.5; specimen col- 
lected by Alfredo Borelli, whose San Francisco = Vil- 
la Monies fide Paynter, 1992). 

PARAGUAY: [Depto. Caaguazu]: Pastoreo [ap- 
proximately 25°23'S, 55°52'W] (MCZ 34214-15). 
[Depto. Central]: Asuncion [25°16'S, 57°40'W] 
(FMNH 9496). Colonia Nueva Italia [25°37'S, 
57°30'W] (FMNH 42281). 

Stenocercus erythrogaster 

COLOMBIA: [Depto. Magdalena]: Rio Frio 
[30-450 m; 10°55'N, 74°10'W] (MCZ 29707). Santa 
Malta Mountains [approximately 10°50'N, 73°40'W] 
(MCZ 11303). Rio Toribio, Hacienda "Papare," sec- 
ond river on road from Cienaga to Santa Marta 
[11°03'N, 74°14'W] (FMNH 165153). [Depto. San- 
tander]: San Gil [1,095 m; 06°33'N, 73°08'W] 
(ANSP 24136, MCZ 36877). 



Stiglich (1922) gives no further information about its 
location. 

-' The specimen was collected in November or De- 
cember 1913 by Jose Steinbach, who collected at a 
locality known as "Las Juntas" during that same pe- 
riod (Paynter, 1992). The two locahties are assumed 
to be the same. The variant spelling "Yuntas" does 
not appear in any sources consulted. 



Stenocercus fimbriatus 

PERU: No specific locality: (FMNH 56070). 
Depto. Huanuco: ca. 35 km NE Tingo Maria, Hcda. 
Santa Elena, ca. 1000 m [approximately 08°57'S, 
76°02'W] (LSUMNS 26966-67). Approximately Vi 
mile E Universidad Agraria de La Selva, Tingo Maria, 
vicinity of Rio Huallaga [3, 09°18'S, 75°59'W], 
USNM 193684. [Depto. La Lihertad]: E Peru, La 
Pinita, Rio Mixiolla, tributaiy of upper [Rio] Hualla- 
ga, 3500 ft [1,067 m; approximately 08°16'S, 76°58'W; 
see footnote 20] (AMNH 56797-98). [Depto. Lor- 
ETO]: E Peru, Contamana, Ucayali River valley [134 
m; 07°15'S, 74°54'W] (AMNH 56803). E Peru, E of 
Contamana on trail to Contaya, 700 ft [213 m; ap- 
proximately 07°15'S, 74°54'W] (AMNH 56781-82). E 
Peru, Pampa Hermosa, mouth of Rio Cushabatay, 
500 ft [152 m] [1, 07°12'S, 75°17'W] (AMNH 56788, 
56790-92, 56794-96, 56801-02). Mishana, Rio Na- 
nay, Estacion Biologica Cauicebus, 150 m [03°53'S, 
73°27'W] (USNM "222377). Mishuana [ = Mishana; 
150 m, 03°53'S, 73°27'W] (KU 212628). Depto. Ma- 
DREDE Dios: Pakitza Station [Rio Manu], Manu Na- 
tional Park [4, 11°56'S, 71°17'W] (Victor R. Morales 
18235). Depto. Ucayali: Rio Curanja, Balta, ap- 
proximately 300 m [approximately 10°08'S, 71°13'W] 
(LSUMNS 17519, 25402-04, 26720-23). Alto [Rio] 
Purris, Alto [Rfo] Curanja, Igarape Champuiaco 
[9°34'S, 70°36'W] (MCZ 61226)" Peru/Brazil frontier, 
Utoquinia Region, 1000 ft. [305 m; approximately 
08°00'S, 74°00'W]22 (AMNH 56789, 56799-800). 

Stenocercus scapulari^^ 

PERU: No specific locality: (FMNH 56444). 
[Depto. Junin]: Chanchamayo, 1200 m [approxi- 
mately 11°03'S, 75°47'W] (FMNH 40608-11). Pere- 
ne, 1200 m [10°58'S, 75°13'W] (MCZ 49580-81). 
Tarma, Chanchamayo, 1300 m [11°25'S, 75°42'W] 
(FMNH 45522). [Depto. Puno]: Sagrario, Rio Qui- 
tun [approximately 1,020 m; 13°55'S, 69°41'W] 



-- The region referred to is north to northeast of 
Pucallpa. The variant spellings Utoquinia, Utoquinea, 
and Uroquinea are in the literature and are applied 
to a right-bank tributary of the Rio Ucayali, a village 
on the Rio Ucayali, and an airstrip on the Rio Uto- 
quinia near the Brazilian border. The entire region is 
less than 500 m in elevation except for a small raised 
area near the Brazilian border that attains nearly 800 
m and that is apparently the source of the Rio Uto- 
quinia. 

^^ The occurrence of Stenocercus scapularis at Rur- 
renabaque. El Beni Department, Bolivia, as reported 
for two specimens in the AMNH (Burt and Burt, 
1931: 273) is apparently based on a misidentification. 
These specimens are probably either S. prionotus 
(most likely) or S. caducus (see Distribution Patterns 
in Stenocercus prionotus). 



218 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3 



(FMNH 40408). "Camp 4" [between Santo Domingo 
and La Pampa; approximately 13°44'S, 69°37'W]-^ 
(FMNH 40409). Juliaca, Lake Aracona, 16,600 ft. 
[shipping point only; correct locality is on the right 
laank of the Rio Inambari, 1,830 m, 13°30'S, 
70°00'W]^'^ (AMNH 1701). 



-^ According to the field catalogue in the FMNH 
Mammal Di\dsion the collector, Colin Sanborn, was 
in Santo Domingo on 20 October 1941 and in La 
Pampa on 23 October (see also notes in Stephens and 
Traylor, 1983). The specimen FMNH 40409 was col- 
lected 21 October, and thus "Camp 4" is assumed to 
be between these points. 

-''The specimen was collected in 1900 by H. H. 
Keays, who collected many mammals and other ver- 
tebrates in southern Peru, primarily for the American 
Museum of Natural History. It is clear that most of 
the specimens labeled with the locality "Juliaca" (a 
town on the Peruvian altiplano near Lake Titicaca) 
actually came from farther north in the Rio Inambari 
valley. Allen (1900: 219; 1901: 41) provides the fol- 
lowing information: 

The Museum has recently received two small col- 
lections of mammals made by Mr. H. H. Keays, at 
Juliaca, in southeatern Peru, a little to the west- 
ward of Lake Titicaca. Mr Keays writes: "Our 
camp is situated in the loop of the Inambaiy River. 
The countiy is very broken, with deep narrow can- 
ons, and is covered with a dense undergrowth oi 
shrubs and vines, with here or there a palmetto or 
a cedar rising above the surrounding vegetation." 
He gives the" altitude as 6000 feet [1,830 m], and 
the position as latitude 13°30' S., longitude 70° W. 

... it is necessary to correct a misleading statement 
in my former paper in respect to the locality where 
the . . . collections were made. Mr. Keays's post- 
office address was Juliaca, and through lack of ex- 
plicit information, it was inferred that the Inca 
Mines, where he collected, were in the immediate 
vicinity of Juhaca . . . the Inca Mines are situated 
about 200 miles northeast of JuHaca, on the east 
side of the Andes, on the Inambaiy River, a trib- 
utaiy of the Amazon, and at a much lower altitude 
than Juliaca. The altitude and geographical position 
were correctly given in the former paper, but in 
place of Juliaca, . . . read Inca Mines. 

Keays's information quoted by Allen places the local- 
ity on the right bank of the Rio Inambari in the foot- 
hills of an outlying Andean spur separating the Rio 
Inambari from upper tributaries of the Rio Tambo- 
pata. I have not located a Lake Aracona and suspect 
that this is an error for Lake Aricoma, a high Andean 
lake on tlie route between Juliaca and the location of 
Keays's camp. However, it is not at all clear why this 
name is associated with the locality. No notes or cor- 
respondence of Keays are in the AMNH mammal 
department archives for further clarification (R. S. 



Stenocercus sp. 

PERU: Depto. Amazonas: 17 km ENE Balsas 
[06°49'S, 7S°00'W] (ROM 16458). 

LITERATURE CITED 

Allen, J. A. 1900. On mammals collected in south- 
eastern Peru, by Mr. H. H. Keays, with descrip- 
tions of new species. Bulletin of the American 
Museum of Natural History, 13: 219-227. 

. 1901. On a further collection of mammals 

from southeastern Peru, collected by Mr. H. H. 
Keays, with descriptions of new species. Bulletin 
of the American Museum of Natural Histoiy, 14: 
41-46. 

Aquino, A. L., N. J. Scott, and M. Motte. 1996. 
Lista de anfibios y reptiles del Museo Nacional 
de Historia Natural del Paraguay (Marzo, 1980 — 
Setiembre, 1995), pp. 331^00. In O. R. Marti- 
nez (ed.), Colecciones de Flora y Fauna del Mu- 
seo Nacional de Historia Natural del Paraguay. 
San Lorenzo, Paraguay: Museo Nacional de His- 
toria Natural del Paraguay. 

AviLA-PlRES, T C. S. 1995. Lizards of Brazilian Ama- 
zonia (Reptilia: Squamata). Zoologische Verhan- 
delingen, Leiden, 299: 1-706. 

Balzan, L. 1931. Viaggio di Esplorazione nolle Re- 
gioni Centrali del Sud America. Milan, Italy: Fra- 
telli Treves, xix + 368 pp. 

BOULENGER, G. A. 1898. A Hst of the reptiles and 
batrachians collected by the late Prof L. Balzan 
in Bolivia. Annali Museo Civil Storia Naturali di 
Geneva, series 2, 19: 128-133. 

. 1901. Further descriptions of new reptiles 

collected by Mr P. O. Simons in Peru and Bo- 
livia. Annals and Magazine of Natural History, 
series 7, 7(42): 546-549. 

1903. List of the batrachians and reptiles col- 



lected by M. A. Robert at Chapada, Matto Gros- 
so, and presented by Mrs. Percy Sladen to the 
British Museum (Percy Sladen Expedition to 
Central Brazil). Proceedings of the Zoological 
Society of London, 1903, II(I): 69-70. 

Brown, F. M. 1941. A gazetteer of entomological sta- 
tions in Ecuador. Annals of the Entomological 
Society of America, 34(4): 809-851. 

Burt, C. E., and M. D. Burt. 1931. South American 
lizards in the collection of the American Muse- 
um of Natural Histoiy. Bulletin of the American 
Museum of Natural History, 61: 227-395. 

Cadle, J. E. 1991. Systematics of lizards of the genus 
Stenocercus (Iguania: Tropiduridae) from north- 
ern Peru: new species and comments on rela- 
tionships and distribution patterns. Proceedings 



Voss, personal communication). Stephens and Traylor 
(1983) discuss "Inca Mines" under their entry for 
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New Species of Stenocercus from Peru and Bolivia • Cadle 



219 



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OF THE 



seum 



(US ISSN 0027-4100) 



On the Subfamily Xylophagainae 
(Family Pholadidae, Bivalvia, Mollusca) 



RUTH D.TURNER 



MCZ 
'.IBRARY 

JAN 2 7 2003 



HARVARD 
UNIVERSITY 



HARVARD university 

CAMBRIDGE, MASSACHUSETTS, U.S.A. 



VOLUME 157, NUMBER 4 
31 October 2002 



(US ISSN 0027-4100) 



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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 
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. 

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© The President and Fellows of Harvard College 2002. 



ON THE SUBFAMILY XYLOPHAGAINAE 
(FAMILY PHOLADIDAE, BIVALVIA, MOLLUSCA) 



RUTH D. TURNERS 

CONTENTS 



Editorial Note 223 

Abstract 223 

Introduction 224 

Species Groups in the Genus Xylophaga 225 

Variation 227 

Preliminaiy Report on the Results of 

Experiments on the Ecology of Deep- 
Sea Wood Borers and the Role of Wood 

in the Deep Sea 234 

Systematic Account 236 

Genus Xylophaga Turton 1822 236 

Xylophaga concava Knudsen 236 

Xylophaga gerda Turner new species 237 

Xylophaga grevei Knudsen 238 

Xylophaga clenchi Turner and Culliney ... 239 

Xylophaga supplicata Tald and Habe 241 

Xylophaga whoi Turner new species 242 

Xylophaga profunda Turner new species 243 

Xylophaga ahyssonuu Dall 245 

Xylophaga dupUcata Knudsen 247 

Xylophaga muraokai Turner new species 247 

Xylophaga atlantica Richards 249 

Xylophaga washingtona Bartsch 250 

Xylophaga rikuzenica Tald and Habe 252 

Xylophaga depalmai Turner new species 253 

Xylophaga guineensis Knudsen 256 

Xylophaga niexicana Dall 257 

Xylophaga tipperi Turner new species 259 

Xylophaga hayeri Turner new species 260 

Xylophaga japonica Tald and Habe 261 

Genus Xylopholas Turner 1972 262 

Xylopholas altenai Turner 263 

Genus Xyloredo Turner 1972 264 

Xyloredo nooi Turner 265 

Xyloredo ingolfia Turner 266 

Xyloredo naceli Turner 267 

Acknowledgment 268 

Literature Cited 268 



' Department of Mollusks, Museum of Compara- 
tive Zoology, Harvard University, Cambridge, Mas- 
sachusetts 02138. 



Editorial Note. Professor Emerita Ruth Dixon 
Turner died on 30 April 2000 and was for the last 
several months of her life severely disabled; in fact, 
her active work as a researcher was considerably fore- 
shortened by medical problems beginning in about 
1995. Among her Nachgelassene Werke was an im- 
portant manuscript on the systematics of the deep- 
sea pholadid bivalve genera Xylophaga, Xyloredo, and 
Xylopholas, a manuscript that she had been preparing 
for a number of years and one that had the active 
support of the U.S. Department of Defense's, then. 
Office of Naval Research (ONR). Professor Turner 
was unquestionably a leading world authority on 
these taxa and had posted this document, in its pre- 
liminary draft form, on a Web site; after her retire- 
ment and the beginning of the illnesses that plagued 
her, the manuscript was removed from the Web site 
witli tlie mtent of readying it for formal publication. Two 
outside authorities. Dr. Jorgen Knudsen of the Zool- 
ogisk Museum, K0benhavns Universitet, K0benhavn, 
Denmark, and Dr. K. Elaine Hoagland, then at the 
Association for Systematic Collections, Washington, 
D.C., were solicited to make criticisms, and these, 
along with my own, were incoi-porated into a more 
advanced revision of the text prepared by Ms. Helene 
Ferranti, a long time coworker and associate of Pro- 
fessor Turner. Ms. Ferranti agreed to revise this 
Nachlass in accordance with the comments of the re- 
viewers and to update its content and organization. 
Having collaborated with Professor Turner on the 
subject of deep-sea bivalves, Ms. Ferranti is credited 
herein as the person responsible for the final com- 
pletion and revision of this valuable text. The new 
species described, for which specimens are available 
for study in the Museum of Comparative Zoology 
(MCZ), and the taxonomic suggestions incorporated 
into the text are to be credited to Professor Turner. 

Kenneth J. Boss 
Editor 

Abstract. The provisional grouping of the species 
of the bivalve genus Xylophaga suggested by Turner 
and Culliney is further elaborated, with 37 species 
assigned to six groups depending on characteristics of 



Bull. Mus. Comp. ZooL, 157(4): 223-307, October, 2002 223 



224 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



the mesoplax, siphons, muscle scars, and method of 
reproduction. Three cases of variation in species of 
Xylophaga are discussed: variation in response to dif- 
ferent substrates, variation that possibly is genetic, 
and variation in a normal growth series. Some obser- 
vations are made regarding the ecology of deep-sea 
wood borers based on experiments carried out with 
wood panels; these support the hypothesis that wood 
contributes to the growth and diversity of deep-sea 
organisms and that the Xylophagainae contribute sig- 
nificandy to the food chain by converting wood to a 
usable form. A section on systematics considers 19 
species of Xylophaga, of which 7 are new, as well as 
the monospecific genus Xylopholas and three species 
of Xyloredo. Detailed descriptions are given of new 
species. 

INTRODUCTION 

The Xylophagainae, a subfamily of the 
Pholadidae, is composed of the genera Xy- 
lophaga Turton, 1822, Xyloredo Turner, 
1972, and Xylopholas Turner, 1972. Species 
in the genus Xijlophaga range in depth 
from just below low tide (X dorsalis Tur- 
ton) to depths of 7,000 m (X grevei Knud- 
son), whereas species in Xyloredo and Xy- 
lopholas are restricted to the deep sea 
(depths of 239 to more than 2,000 m). 
Species in Xyloredo range in depth from 
1,737 to more than 2,000 m (X nooi Turn- 
er, 1972) and species in Xylopholas range 
from 239 to 366 m (X altenai Turner, 
1972), with one lot dredged in 2,550 m off 
Port Victoria, Sao Tome, Gulf of Guinea. 
However, the Sao Tome specimens were 
boring in coconut shells and may not have 
been living at that depth. 

The Xylophagainae are marine, cosmo- 
politan, and range from moderate to abys- 
sal depths. All of the Xylophagainae, so far 
as known, are wood borers, and all are 
sublittoral. Only in high latitudes do they 
compete with shipworms (Teredinidae) in 
cold boreal waters. So far as known, the 
Xylophagainae do not occur intertidally, or 
in floating wood. Wood containing speci- 
mens of the Xylophagainae usually has 
been obtained by dredging. Occasionally, 
specimens may be obtained from water- 
logged wood that has been brought up and 
thrown ashore during a storm after being 
on the bottom for some time. 



Until recently, the Xylophagainae were 
considered to be deep-sea organisms of lit: 
tie or no economic importance. They were 
rare curiosities, of interest mostly in their" 
role of recycling wood on the continental 
shelf and slope and in the abyss, largely 
beyond the depth range of the teredinids. 
The Xylophagainae often were referred to 
as shipworms and because of the ephem- 
eral, patchy distribution of wood in the 
deep sea were thought to have little im- 
pact on ecological processes. With the ex- 
tension of human activities into the deep 
sea for fishing (especially trap fishing for 
lobsters and crabs), as well as for archae- 
ology, mining, monitoring currents, and 
other activities, these borers are now gen- 
erally considered to be pests. 

Species in the genus Xylophaga (Xylo = 
wood, phaga = eating) are restricted to 
wood, woody plants, and structures made 
of wood found in the deep sea. In com- 
mon with the teredinids (shipworms), they 
have symbiotic bacteria in their gills (Wa- 
terbuiy et al., 1983). These bacteria are be- 
lieved to have the ability to digest cellulose 
and probably to fix nitrogen. Collaborative 
work with Dr. Waterbury, microbiologist at 
Woods Hole Oceanographic Institute, was 
unable to culture cellulose-digesting bac- 
teria from the gills of X atlantica, but ev- 
idence was found of cellulose enzymes in 
the gill tissue. 

The three genera of Xylophagainae may 
be briefly characterized as follows: 

Genus Xylophaga Turton. Siphons rela- 
tively short, of equal length or with the 
excurrent siphon truncated, and often 
capable of retraction between the 
valves. Burrow seldom more than five 
times the length of the valves and often 
with a chimney of fecal pellets lining the , 
posterior end of the burrow. | 

Genus Xylopholas Turner. Shell typical 
but with the animal extended and with 
lateral plates on the siphons. 

Genus Xyloredo Turner. Shell typical but 
animal elongated and producing a tere- j 
dinidlike burrow that is lined with a cal- 



Xylophagainae • Turner 



225 



careous tube marked with distinct 
growth rings and margined anteriorly 
with a periostracal band. 

The Xylophagainae are often confused 
with teredinids, but the gills and digestive 
and reproductive organs in the Xylopha- 
gainae do not extend posteriorly beyond 
the valves. In addition, the Xylophagainae 
do not have pallets to close the entrance 
to their burrows or apophyses for the at- 
tachment of the foot muscles. In common 
with the teredinids, but unlike species in 
the pholadid genera Martesia and Ligno- 
pholas, the only other genera of wood-bor- 
ing pholadids, the Xylophagainae have a 
large wood-storing cecum and probably 
utilize the wood in which they bore as 
food. For details of anatomy, see Purchon 
(1941) for Xijlophaga dorsalis Turton, and 
Turner (1955) for X atlantica. 

SPECIES GROUPS IN THE 
GENUS XYLOPHAGA 

Genus Xijlophaga Turton. Xylophaga Turton, 1822, 
Conchylia Insularum Britanicarum, p. 253 (type 
species, Teredo dorsalis Turton, 1819). 

Species in this genus are characterized 
by teredolike shells that lack apophyses 
and have a divided mesoplax that is vari- 
able in shape and size. A chrondrophore 
and internal ligament are present. The si- 
phons are variable, united for part or all of 
their length, with the excurrent siphon of- 
ten truncated. The visceral mass and gills 
do not extend beyond the valves posteri- 
orly. The wood-storing cecum is large. 

In his discussion of the taxonomy of Xy- 
lophaga, Knudsen (1961) believed that the 
use of subgenera was not feasible and 
would only lead to the creation of a large 
number of monotypic subgenera that 
would be of limited value. This is still par- 
tially true but new species described in 
this report and the additional material now 
available concerning other species have 
made possible a provisional grouping of 
the species, as suggested by Turner and 
Culliney (1971). See also Hoagland and 



Turner (1981) and Hoagland (1983). This 
grouping is helpful when discussing rela- 
tionships and geographic distribution. The 
characters used for grouping the species 
are those mainly of the mesoplax and si- 
phons in conjunction with the muscle scars 
and methods of reproduction (see Text- 
Fig. 1). The mesoplax is a transverse plate, 
usually wider than long, that straddles the 
valves at the umbos and partially or com- 
pletely covers the posterior end of the an- 
terior adductor muscle. The mesoplax may 
be composed of one or two parts. The im- 
portant character of the mesoplax is the 
presence or absence of a ventral portion 
and tubes; the more detailed characters, 
such as the presence of lobes, seem to be 
of specific value only. Siphonal characters 
include the relative length of the two si- 
phons, the presence or absence of cirri at 
the apertures, and the type of siphonal 
folds, which may or may not have lappets 
or fringes. 

Not all characters are known for all spe- 
cies and a few species seem to be transi- 
tional between groups. There is no ques- 
tion that more material is needed before 
definite statements can be made concern- 
ing the formal use of subgenera. However, 
the grouping of species as presented here 
does offer an opportunity to speculate on 
the possible origin and evolution of the ge- 
nus and to focus attention on the types of 
information that should be considered in 
future studies. Comparative anatomical 
and molecular studies are greatly needed 
but it probably will be some time before 
these can be completed because well-pre- 
served specimens of deep-sea Xylophaga 
are rare and difficult to obtain. 

If we consider species with simple si- 
phons of equal length and a mesoplax of 
two simple flat to slightly curved plates to 
be the basic type, it is possible to group 
the species in what appears to be a devel- 
opmental series of six groups. This list 
does not include all nominal Xylophaga. 
The groups may be characterized as fol- 
lows: 



226 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



siphonal retractor 
pedal retractor 
posterior adductor 



umbonal reflection 
anterior adductor 




ventral adductor 
1 bona! -ventral ridg 



Text-Figure 1 . Nomenclature of parts of Xylophaga. (1) Internal view of left valve showing relative position of muscle scars. (2) 
External view of left valve. (3) Dorsal view of animal with siphons retracted. (4) Lateral view of entire animal with siphons 
extended. 



Group 1. Mesoplax composed of two 
simple flat or slightly cuived plates lo- 
cated posterior to the anterior adductor 
muscle and standing erect. Siphons of 
equal length or with the excurrent si- 
phon slightly shorter, cirri on one or 
both siphons present or absent. Group 
1 includes X. erecta Knudsen, X. con- 
cava Knudsen, and X. gerda Turner n. 
sp. 

Group 2. Mesoplax composed of two 
plates that may be curved, flat, sculp- 
tured, or smooth, set at an acute angle 
to each other, lack dorsal tubes and a 
ventral portion, but cover the anterior 
adductor muscle dorsally. Siphons of the 
same length or with the excurrent si- 
phon slightly shorter and with large cirri 
on the sides of the excurrent siphon and 
small cirri at the incurrent siphon. 
Group 2 includes X. grevei Knudsen, X. 
wolffi Knudsen, X. hadalis Knudsen, X. 
galatheae Knudsen, X. murraiji Knud- 
sen, X. africana Knudsen, X. pananien- 
.sis Knudsen, and X. clenchi Turner and 
Culliney. 



Group 3. Mesoplax composed of two 
nearly flat plates set at an acute angle to 
each other forming an inverted V, with 
tubes extending from the posterior dor- 
sal surface or longitudinally folded with 
anterior lobes or pores. Mesoplax lack- 
ing a ventral portion and set in a tentlike 
fashion over the anterior adductor mus- 
cle. Siphons nearly the same length and I 
usually with small cirri on both open- 
ings. Group 3 includes X. supplicata 
Taki and Habe, X. lohata Knudsen, X. 
tuhidata Knudsen, X. bniuni Knudsen, 
X. obtusata Knudsen, X. whoi Turner n. 
sp., and X. profunda Turner n. sp. 

Group 4. Mesoplax composed of two 
plates that have a small to large ventral 
portion, the dorsal portion being 
smooth, folded, or lobed. Siphons of the 
same length or with the excurrent si- 
phon slightly shorter and with cirri or 
papillae at one or both openings. Group 
4 includes X. abyssorum Dall, X. dupli- 
cata Knudsen, X muraokai Turner n. 
sp., X. foliata Knudsen, and X. atlantica 
Richards. 



Xylophagainae • Turner 



227 



Group 5. Mesoplax composed of two 
plates that are more or less triangular in 
outline, with a ventral portion ranging 
froin very narrow to more than one half 
the width of the dorsal portion. The ex- 
current siphon may vary in length from 
one half to three quarters that of the 
incurrent siphon and have cirri, or it 
may be truncated just posterior to the 
valves and have dorsal lobes or folds ex- 
tending from the truncation along the 
dorsal surface of the incurrent siphon 
for part or all of its length. Group 5 in- 
cludes X. washingtona Bartsch, X ri- 
kuzenica Taki and Habe, X. aurita 
Knudsen, X. turnerae Knudsen, and X. 
praestans E. A. Smith. 

Group 6. Mesoplax composed of two 
more or less ear-shaped plates some- 
what coiled posteriorly. Excurrent si- 
phon truncated near the posterior end 
of the valves, continuing as lateral lobes 
extending from the truncation along the 
dorsal surface of the incurrent siphon. 
These lobes may vary in width but are 
always fringed. Group 6 includes X. dor- 
salis Turton, X. depahnai Turner n. sp., 
X. guineensis Knudsen, X. mexicana 
Dall, X. tipperi Turner n. sp., X. baijeri 
Turner n. sp., X. globosa Sowerby, X.ja- 
ponica Taki and Habe, and X. indica 
Smith. 

Most species are known only from the type 
series and these have all been studied by 
the author except X. indica, and the spe- 
cies described by Taki and Habe. Howev- 
er, paratype specimens received through 
the kindness of Dr. Habe are in the col- 
lection of the MCZ. They include X. ja- 
ponica, X. rikuzenica, X. teramachi, and X. 
supplicata, although unfortunately all lack 
the mesoplax except the last. Two species, 
X. teramachi Taki and Habe (Taki and 
Habe, 1950) and X. tomlini Prashad (Pras- 
had, 1932); are known only from the valves 
and remain unassigned. A map showing 
the distribution of species of Xylophaga is 
provided in Text-Figure 2. 

Nineteen of the 37 species oi Xylophaga 



listed in the groups above as well as the 
monospecific genus Xylopholas and the 
three species of Xyloredo are considered 
in the section on systematics. Some spe- 
cies are discussed more fully than others 
but the distinctive characters have been 
given for all. For example, Xylophaga mex- 
icana Dall and Xijlophaga abyssorum Dall 
are fully described because these names 
were based on valves only and were vir- 
tually nomina dubia. By matching the 
valves of the holotypes with complete 
specimens, it has been possible to fix the 
names of these species. If additional char- 
acters or records are given for a well-de- 
scribed species, a reference is made to the 
original description. Detailed descriptions 
are given for new species. 

VARIATION 

Knudsen (1961) aptly stated that very 
little was known about variation in species 
of Xylophaga. Unfortunately, large series 
of any one species seldom have been avail- 
able for study because inaterial usually is 
obtained from small pieces of wood or oth- 
er plant material that has been dredged or 
occasionally thrown ashore as driftwood. 
Of the 30 species listed by Knudsen and 
the 7 species described as new in this pa- 
per, 24 are known from fewer than 10 
specimens; only 7 species are known from 
series of more than 100 specimens. Most 
are known from only one or two localities 
and often all specimens are froin a single 
piece of wood, and may all be of the same 
set, that is, have settled at the same time. 
Consequently, it is not surprising that all 
specimens in any one lot are quite similar. 
Only since the beginning of deep-sea test- 
ing and the use of the submersible DSV 
Alvin to place experimental wood islands 
at great depths has it been possible to ob- 
tain sufficient material to study intraspe- 
cific variation in this subfamily. 

The earliest work of this sort was done 
by the U.S. Naval Civil Engineering Lab- 
oratory (USNCEL) and the Navy Ocean- 
ographic Office (NOO). Three cases of 
variation based on this material are re- 



228 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 




Text-Figure 2. Distribution map of species of Xylophaga. 



ported here. The first case involves varia- 
tion in response to different substrata, the 
second case exliibits variation that possibly 
is genetic because the substrate and all 
other parameters were as nearly uniforin 
as possible, and the third case illustrates 
variation in a normal growth series. 

Variation Due to Different Substrates. 
Variation in Xylophaga mashingtona in re- 
sponse to the substrate can be deinonstrat- 
ed with material from the USNCEL tests. 
A series of 10 panels of different types of 
wood were attached to a submersible test 
unit (STU) that was submerged from April 
1965 to May 1966 off San Miguel Island, 
California (34°06'N, 120°42'W) at 2,370 ft 
(730 m) (see Table 1 and Text- Figs. 3-9). 
The 2 X 6 X 0.5-in (50.8 X 152.4 X 12.7- 
min) wood panels were all attached to the 
same rack on the STU so that they would 
be resting just above the mudline. Con- 
sequently, all factors affecting the borers 



were as nearly identical as possible except 
the substrate (i.e., the species of wood) on i 
which the borer larvae settled and into j 
which they would bore. Text-Figures 3—8 
illustrate typical specimens from each of 
the wood panels; Text-Figure 9 shows 
specimens from a phenolic laminated rod. 
It is interesting to note that the dorsal 
plates in all specimens are remarkably uni- 
form, varying only slightly in length/width 
proportions. Even the specimens taken 
from the phenolic laminated rod could be 
identified by the dorsal plates. 

The general shape of the valves with the 
high posterior slope also remained rather 
constant except in the extremely steno- 
morphic (stunted) specimens from Afam- 
beaii and the phenolic laminated rod. It is 
difficult to explain the proportionate size 
of the larval valves on specimens boring 
into harder materials except that these 
speciinens had not greatly increased in di- 



Xylophagainae • Turner 



229 



Table 1. Variation of Xylophaga washingtona burrows in different types of wood. 



Wood 



No. 


Burrow 


Burrow 


Specimens 
Examined 


Lens^th 


Diameter 


(mm) 


(mm) 



Remarks 



Cedar 

Ash 

Maple 

Pine 
Oak 

Fir 

Redwood 

Greenheart 

Afambeou 

Antidesma pulvinatum 



125 


21.0 


5.0 


75 


11.0 


5.5 


35 


15.0 


5.0 


±50 


15.0 


6.0 



50 25.0 4.5 many dead, often three or four specimens in one 

enlarged cavity where burrows ran together. 
Heavily attacked, particularly at one end. 

many dead, burrows running together, panel 
heavily attacked, particularly at one end. 

well distributed, with a little more concentration 
around the hole at one end. 

clustered at one end, newly settled to adult. 

evenly distributed, shells yellowish-green from 
wood. 

46 12.0 4.5 many newly settled, often cut into burrows of oth- 

er specimens. 

specimens stained dark red brown by wood. 

concentrated around edges. 

concentrated at one end, all very small, many in 
umbo stage veliger, 15 small depressions with- 
out animals. 
150 0.10 0.05 many newly settled, just beginning metamorpho- 



46 


14.0 


4.5 


75 


2.75 


1.75 


19 


0.75 


0.05 



Cedar 




5 mm 



Pine 




5 mm 




I 1 

1 mm 



1 mm 



Text-Figure 3. Typical specimens of Xylophaga washingtona collected from submerged cedar (top) and pine (bottom). 



230 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Ash 




5 mm 
Text-Figure 4. Typical specimens of Xylophaga washingtona collected from submerged ash (top) and maple (bottom). 



Oak 




5 mm 



Fir 




5 mm 



Text-Figure 5. Typical specimens of Xylophaga washingtona collected from submerged oak (top) and fir (bottom). 



Xyloph AGAIN AE • Turner 231 



Redwood 



Greenheart 




Text-Figure 6. Typical specimens of Xylophaga washingtona collected from submerged redwood (top) and greenheart (bottom). 




Afambeau 



0.1 mm 
Text-Figure 7. Typical specimens of Xylophaga washingtona collected from a submerged panel of Afambeau. 



232 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 




Antidesma pulvinatum 



Text-Figure 8. Typical specimens of Xylophaga washingtona collected from a submerged panel of Antidesma pulvinatum. 



ameter as they bored so that the lai"val 
valves were not inturned with the growth 
of the umbos. Evidently little or no attri- 
tion had occurred at the umbonal area or 
the embryonic valves would have been 
worn away. 

The most variable characters in this se- 
ries are the size of the valves and the num- 
ber of denticulated ridges on the anterior 
slope in relation to the length of the valves. 
Specimens from cedar, pine, ash, maple, 
oak, and fir (Text-Figs. 3-5) all were drawn 
to the same scale, as shown by the 5-miTi 
scale bar, and are arranged in order of de- 
creasing size. The nuiTiber of denticulated 
ridges on the anterior slope of these spec- 
iiTiens varies from 12 to 20. Specimens 
froin redwood, greenheart, Afambeau, and 
Antidesma pulvinatum (Text-Figs. 6—8) 
"were drawii to their own scales and, al- 
though these speciinens are much siTialler, 
they have as many or more ridges on the 
anterior slope. The specimen removed 



from the phenolic laminated rod (Text-Fig. 
9) has 44 denticulated ridges. Correlated 
with the increased hardness of the sub- 
strate and the additional denticulated ridg- 
es is a proportional increase in the size of 
the posterior adductor muscle and its scar. 
This suggests the enlargement of the pos- 
terior adductor inuscle in response to in- 
creased activity of boring. The general 
shape and sculpturing of the inuscle scar 
was similar in all specimens. No noticeable 
variation was found in the siphons, except 
size, regardless of the substrate. However, 
specimens that were able to bore deeply 
into the wood usually formed a chimney 
composed of compacted fecal inaterial lin- 
ing the posterior end of the burrow. 

Two periods of settlement apparently 
occurred on some of the panels, because 
specimens of two age groups could be 
found. However, it is impossible to say 
whether the specimens removed from the 
Antidesma were of the second set or if 




Xylophagainae • Turner 



233 




1 mm 



Phenolic Laminated Rod 

Text-Figure 9. Typical specimens of Xylophaga washingtona collected from a submerged phenolic rod. 



they had simply been unable to increase 
in size because of the hardness, chemical 
composition, or both of the wood. Because 
the specimens apparently were alive at the 
time the wood was reinoved from the wa- 
ter and because only a few rows of dentic- 
ulated ridges were present, the inference 
was made that these specimens probably 
belonged to a second set. Certainly the lar- 
val shells shown in Text- Figure 8 must be 
from a second set. The number of speci- 
mens examined and the maximum length 
and diameter of the burrows for each type 
of wood are given in Table 1 . 

Variation in the Mesoplax o/ Xylophaga 
depalmai Turner n. sp. Approximately 300 
specimens of X. depalmai n. sp. were ob- 
tained from tests conducted by the NOO 
about 2—3 miles east of Fort Lauderdale, 
Florida (26°04'N, 80°04'W) in depths 
from 100 to 500 ft (30.5 to 152.5 m) (see 
Table 14 under the description of X de- 
palmai n. sp. for information giving panel 
numbers, depth, and dates of exposure). 
In this species, the general shape of the 
valves, the siphons, and the muscle scars 
show little variation but the mesoplax is 
extremely variable. The mesoplax is typi- 
cally bilaterally symmetrical, ear-shaped, 
longer than wide, with the two halves 
coiled inward at the posterior end, and 



with a long medial line where the two 
halves meet (Plate 24, Figs. 16, 17). In nu- 
merous specimens, the mesoplax was not 
bilaterally symmetrical, but one half was 
considerably shorter than the other and of- 
ten appeared malformed (Plate 24, Figs. 
11—15). In several specimens, the ventral 
surface of the two halves of the mesoplax 
was fused by the periostracal covering, al- 
though the dorsal surface still appeared di- 
vided. In other specimens, the mesoplax 
was elongated, the coiled posterior ends, 
instead of curling inward toward each oth- 
er, remained nearly straight or curled 
slightly outward, with the ventral surface 
being completely fused (Plate 24, Figs. 1- 
5). In two specimens, the two halves of the 
mesoplax had completely fused dorsally 
and ventrally, although the lines of fusion 
remained clearly visible. The tapered pos- 
terior end of this cornucopialike mesoplax 
coiled slightly ventral and to the left. 

Such variation is in marked contrast to 
the uniformity seen in the mesoplax of X. 
washingtona Bartsch. It is impossible to 
say whether this variation is genetic or eco- 
logic but we are able to say that all types 
of the mesoplax of X. depalmai were found 
in a single panel retrieved from a depth of 
approximately 300 ft (91.44 m). Variation 
in the mesoplax is a factor that must be 



234 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



taken into consideration when evaluating 
species in this genus. 

Variation Exhibited in Growth Series. 
In most species of Xylophaga, the dorsal 
plate is quite simple and the mesoplax of 
the young is similar to that of the adult, as 
seen for X. washingtona. Dons (1929a,b) 
described briefly and illustrated a similar 
situation in X praestans Smith and X. dor- 
salis Turton. 

Some species have more elaborate dor- 
sal plates and for some of these (i.e., X. 
muraokai Turner n. sp., X. hayeri Turner 
n. sp., and X. abyssorum Dall), it is pos- 
sible to build up what appears to be 
growth series. All begin with a simple pos- 
terior covering to the anterior adductor 
muscle that is difficult to differentiate 
among the species in the young stage. As 
calcification of the periostrascal covering 
progresses, the adult form gradually ap- 
pears. In the three species mentioned, the 
characters of the valves and the siphons of 
the young agree with those of the adult 
specimens. Therefore, although living ma- 
terial has not been available and develop- 
mental studies have not been possible, it 
seems reasonable to consider these as 
growth series. Xylophaga muraokai (Plate 
19, Fig. 3) is relatively simple, with the 
dorsal portion becoming thickened and 
joining laterally with the basal portion 
while the ventral flanges enlarge. In X. 
hayeri (Plate 31, Fig. 3), the broad trans- 
verse dorsal portion becomes very con- 
spicuous and in X. abyssorum (Plate 15, 
Figs. 4, 5), the dorsal incurving of the lat- 
eral arms produce an elaborately lobed 
mesoplax. Nothing is known of the allo- 
metric changes that take place during 
growth of the peculiar dorsal plates of X. 
tiibulata Knudsen, X. bruuni Knudsen, X. 
obtusata Knudsen, and X. whoi Turner n. 
sp. 

Detailed studies of these growth series 
will have to await improved techniques for 
handling living material under deep-sea 
conditions, and the ability to rear deep-sea 
species in the laboratoiy. In the meantime, 
it seems best to consider these forms as 



members of growth series rather than dif- 
ferent species, particularly because the se- 
ries in each case was built up from mate- 
rial taken from a single piece of wood. 

PRELIMINARY REPORT ON THE 
RESULTS OF EXPERIMENTS ON THE 
ECOLOGY OF DEEP-SEA WOOD 
BORERS AND THE ROLE OF WOOD IN 
THE DEEP SEA 

The results of the first exposures of 
wood panels at the Woods Hole Oceano- 
graphic Institution (WHOI)-AZuin per- 
manent bottom station south of Woods 
Hole (39°46'N, 70°41'W, in 1,830 m) were 
reported in 1973 (Turner, 1973). At that 
time, the Xylophagainae were postulated ! 
to be the most important deep-sea organ- 
isms involved in converting woody plant 
material to an available food source for 
other organisms. 

Pursuing this theory and to continue the 
studies of the ecology and life history of 
the Xylophagainae, wood panels were ex- 
posed at the Alvin permanent station in 
the Tongue of the Ocean, Bahama Islands, 
on 19-22 January 1974 {Alvin dives 492, 
493, 494, and 495) at a depth of 2,032 m. 
The first of these panels was picked up on 
7 March 1974, frozen immediately, and re- 
turned to Woods Hole where it was ex- 
amined. Newly settled larvae and meta- 
moi^phosing Xylophaga with one to two 
rows of denticulated ridges were removed 
from the panel. The specimens were ap- 
proximately 300 |JLm in length and the 
greatest penetration was about twice the 
depth of the shell. The debris rings sur- 
rounding the burrows were much coarser 
than those made by teredinids. The distri- 
bution of the entrance holes was some- 
what patchy and varied from 5 to 20 cm^. 
The specimens were too young to identify 
because none of the dorsal plates had been 
formed but examination with scanning 
electron microscopy showed a well-devel- 
oped distinctive sculpture on the larval 
shell. 

This first panel from the Tongue of the 
Ocean established that Xylophaga were 



XYLOPHAGAINAE • Turner 



235 



just beginning to settle on the wood a max- 
imum of 48 days after it was implanted in 
the bottom and that settlement of larvae 
could occur in early March, at least at this 
site. 

Three more panels were removed from 
the Tongue of the Ocean station (Tower 
1 — west arm) on 19 April 1975 during Al- 
vin dive 552. I was an observer on this dive 
and as we approached the panels I noticed 
an increase in the number of shrimp and 
galatheid crabs. The panels had numerous 
crabs crawling all over them. Some of the 
crabs had crawled under the plastic mesh 
bags covering the panels and had grown so 
large they could not escape. (Note: After 
the near loss of the panels at the northern 
station because of the heavy attack of bor- 
ers, the decision was made to put the pan- 
els in plastic mesh bags so that the pieces 
could be retrieved if they began to disin- 
tegrate.) The specimens inside the bag 
were carried to the surface v^dth the pan- 
els. When the panel was disturbed, the 
specimens on the outside of the mesh fell 
off. The largest of the 12 crabs was 43 mm 
in length. The diamond-shaped opening of 
the mesh was 5 X 10 mm. The smallest 
crab ineasured 8 mm in length; others 
measured 40, 33, 32, 30, and 24 mm. It is 
obvious that the crabs were finding suffi- 
cient food either in or on the wood to grow 
at a fairly rapid rate. 

The first young crabs to find the wood 
may have fed on the newly settled Xijlo- 
phaga laiA^ae and this might explain the 
patchy distribution of the borers in the 
panels. However, the larger crabs would 
not have stayed on the wood unless there 
was something for them to eat. The crabs 
had to be under 10 mm in leng-th to gret 
under the mesh and if the larvae were not 
settling until early March it would be at 
least early May before the borers had 
grown sufficiently to be a good food source 
for the crabs. Therefore, I think we can 
postulate that the largest crab measured 
grew at least 33 mm in a period of 10 
months. 

Examination of the panels showed a 



rather heavy attack of three species of Xy- 
lophagainae. These included Xyloredo 
nooi Turner and two Xylophaga species, 
Xylophaga clenchi Turner and Culliney 
and X profunda Turner n. sp. The X nooi 
were typical with valves that reached 5 
mm in length and burrows that were 18— 
22 mm in length. The calcareous lining of 
the burrow of the largest specimen was 13 
mm long and 2.5 mm in diameter at its 
anterior end. The smaller species of Xy- 
lophaga, X. clenchi Turner and Culliney, 
also had been obtained previously from 
wood exposed in the Tongue of the Ocean 
by John DePalma of the NOO. This is a 
fairly small species. The valves were 8—10 
mm in length and several of the specimens 
were canying lai"vae on the umbonal area 
of the valves. The laivae measured 0.2 mm 
in length. The large species o{ Xylophaga, 
X. profunda Turner n. sp., had not been 
seen before. The valves were 14 mm in 
length, and one specimen measured 40 
mm to the tip of the siphons. The burrows 
were 45—50 mm in length. Both X clenchi 
and X profunda lined the posterior end of 
their burrows with consolidated fecal pel- 
lets and the burrows of all dead specimens 
contained one or more specimens of cap- 
itellid worms that were feeding on the pel- 
lets as well as the remains of the Xylopha- 
ga. Often the spaces between the valves of 
the borers were filled with the smaller fe- 
cal pellets of the worms. Breaking these 
balls of pellets apart, I always found one 
or two capitellid worms. In the Xylophaga 
burrows and on the surface of the wood, 
I also found two other polychaete worms. 
One belonged to the family Chiysopetali- 
dae and the other to a family of polynoid 
wonns. 

A preliminaiy examination of the stom- 
ach contents of a broken specimen of a 
galatheid showed that the crab had ingest- 
ed some fine chips of wood because iden- 
tifiable cells remained in the material. 
Consequently, we can postulate that the 
crabs were feeding on the Xylophaga, 
probably dead ones. The tissues of the Xy- 
lophaga are so soft that they are unrec- 



236 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



ognizable in such a preliminary examina- 
tion. The crab's stomach also contained se- 
tae of the chrysopetalid worms, a small 
nematode, and some sponge spicules. The 
chrysopetalids probably were feeding on 
the capitellid worms. 

I think five points are worthy of notice: 
1) Xijlophaga in the Tongue of the Ocean 
as well as at the northern Alvin station are 
growing much faster than one would ex- 
pect. 2) The lai-vae of X profunda n. sp. 
were settling in early March. 3) Xijlophaga 
clenchi broods its young and was carrying 
young in mid-April. 4) Probably the crabs 
and worms were also growing faster than 
normal for the deep sea. 5) A food chain 
based on wood and Xijlophaga was being 
developed. 

This lends support to my hypothesis that 
wood is an important source of enrich- 
ment in the deep sea, that it contributes 
to both the diversity of organisms in a giv- 
en area and to their rate of growth, and 
that the Xylophagainae are the most im- 
portant organisms involved in converting 
the wood to a usable form. To my knowl- 
edge, this is the first documented food 
chain for invertebrates in the deep sea. On 
the basis of these simple experiments, it 
now seems conceivable that the slow 
growth rates usually attributed to deep-sea 
animals may be due to lack of food, at least 
for epifaunal forms, rather than being an 
inherent characteristic of the species in- 
volved. 

SYSTEMATIC ACCOUNT 

GENUS XYLOPHAGA TURTON 1822 

Xytophaga concava Knudsen 
Plate 1 

Xijlophaga concava Knudsen, 1961, Galathea Report, 
5: 167-169, figs. 4, 5 {Galathea, station 726, Gulf 
of Panama [5°49'N, 78°52'W] in 3,270-3,672 m). 
Holotype, Zoological Museum, University of Co- 
penhagen; paratyjae, MCZ 235796. 

Distinctive Characters. Posterior slope 
of valves concave when viewed dorsally 
Mesoplax composed of two rather wide, 
erect, curved plates that extend above the 



umbos. Siphons nearly the saine length, 
excurrent slightly shorter with a few large 
cirri, incurrent siphon with many small cir- 
ri (Plate 1, Figs. 2, 3). Chimney apparently 
lacking, not mentioned by Knudsen and 
not found with the single specimen re- 
ported here. 

Remarks. At the time Knudsen de- 
scribed this species he had 4 specimens 
from Galathea station 726 and 25 speci- 
mens from Galathea station 739. Both of 
these stations were given as in Gulf of Pan- 
ama. However, station 726 is about 95 
miles west of the Gulf of Tibuga, Golom- 
bia, whereas station 739 is about 90 miles 
west of Ensenada Guayabo, Panama. A 
single specimen of X. concava was taken 
by the RA^ Pillshury at station 526. This 
locality is about midway between the two 
Galathea stations. 

The PiUsbury specimen agrees closely 
with the description and figures given by 
Knudsen (1961) except that the incurrent 
siphon has a double row of about 25 small 
cirri around the aperture and the excur- 
rent siphon has 6 large cirri (Plate 1, figs. 
2, 3). Concerning the siphons, Knudsen 
stated that "both openings are at the distal 
end, close together, and around theiu 15- 
16 small cirri are present." In Knudsen's 
illustration, the distal ends of the siphons 
appear to be contracted; neither the two 
openings nor the cirri are apparent. The 
contracted condition of Knudsen's speci- 
mens probably accounts for the differenc- 
es noted here. 

Xijlophaga concava is closely related to 
Xijlophaga gerda Turner n. sp. but differs ; 
in the size of the mesoplax, the type of 
siphonal openings, and the chimney. (See 
Remarks under x. gerda.) Xijlophaga con- 
vava also is related to X. erecta Knudsen 
(1961) from the Sulu Sea. Knudsen (1961) i 
reported that no cirri were visible on the! 
siphons of X. erecta, that the posterior ad- 
ductor scar was much broader in X. con- 
cava, and that the posterior slope of the 
valve viewed dorsally was convex in X. er- 
ecta rather than concave. Knudsen's de- 
scription of X. erecta was based on 20 



Xylophagainae • Turner 



237 



specimens and unfortunately no further 
records have been obtained. 

Range. From off Ensenada Guayabo, 
Panama, south to off the Gulf of Tibuga, 
Colombia, in depths from about 915 to 
3,670 m. 

Specimens Examined. COLOMBIA: Galathea, sta- 
tion 739, Gulf of Panama, about 90 miles W of En- 
senada Guayabo (7°22'N, 79°32'W) in 915-975 m 
(dried specimens); Pillsbunj, station 526, about 110 
miles W of Cabo Marzo (6°53'N, 79°27'W) in 3,193- 
3,211 m; Galathea, station 726, Gulf of Panama 
(5°49'N, 7S°52'W) in 3,270-3,670 m. 

Xylophaga gerda^ Turner new species 
Plates 2, 3 

Holofijpe. MCZ 328378. Paratypes, 
MCZ 316741, 316742. 

Type Locality. Gerda, station 499, about 
3 miles off Southwest Point, Great Baha- 
ma Island, Bahama Islands (26°37'N, 
78°56'W) in 155 fathoms (283.96 m). 

Distinctive Characters. Posterior slope 
of valves concave when viewed dorsally. 
Mesoplax composed of two narrow, erect 
cui-ved plates at the posterior end of the 
anterior adductor muscle. Siphons of 
equal length, with a periostracal sheath 
and four or five cirri surrounding the ap- 
ertures. Chimney composed of fecal ma- 
terial agglutinized to a periostracal base 
(Plate 2, Fig. 7). 

Description. Shell globose, fragile, 
reaching 3.0 mm in length and 2.8 mm in 
height, umbos inflated. Pedal angle 110— 
115°. Anterior slope with up to 45 rather 
evenly spaced denticulated ridges. Umbo- 
nal— ventral sulcus narrow and only slightly 
depressed. Disc and posterior slope sculp- 
tured with fine, incised growth lines. Pos- 
terior slope high, flaring, and somewhat 
ear-shaped. 

Inner surface of valves smooth and glis- 
tening. Umbonal— ventral ridge low and in- 
distinct except near the wide, low ventral 
condyle. Chondrophore and internal liga- 



Table 2. Measurements of Xylophaga gerda. 



Length 

(mm) 


lIHiihl 

(min) 


Location 


1.1 


1.0 


Gerda, station 266 


1,5 


1.4 


Gerda, station 266 


1.8 


2.0 


Gerda, station 266 


2.5 
3.0 


2.3 
2.5 


paratype 
holot)/pe 


3.0 


2.8 


Pillsbunj, station 328 


3.3 


3.0 


Pillsbunj, station 944 


3.8 


3.5 


Pillsbunj, station 944 



- Named for RA^ Gerda, Rosenstiel School of Ma- 
rine and Atmospheric Sciences, University of Miami, 
Miami, Florida, whose station 499 is tiie type locality. 



ment small. Posterior adductor muscle 
scar large with faint transverse impres- 
sions, which are best seen externally on an 
entire specimen. Disc separated from pos- 
terior slope by a shelflike ridge (Plate 3, 
figs. 2—5). Pedal and siphonal retractor 
scars not visible. 

Mesoplax composed of two erect, nar- 
row, curved, slightly calcified plates, locat- 
ed just posterior to the anterior adductor 
muscle and not extending above the um- 
bos. 

Siphons long, probably not capable of 
retraction between the valves, of equal 
length, united nearly to the tip, with a thin 
periostracal sheath. Siphonal apertures of 
about equal size, each with four or five 
comparatively large cirri, which appear as 
a common ring of cirri when the siphons 
are retracted. A browii periostracal cylin- 
der containing fecal material may extend 
nearly one half the length of the excurrent 
siphon (Plate 2, Figs. 1-4). Chimney built 
in sections, composed of fine fecal material 
agglutinized on a periostracal lining with 
"leaves" of periostracum extending to the 
outer surface (Plate 2, Figs. 6, 7). Arrange- 
ment of the gills and labial palps typical 
for the genus, foot large but not muscular, 
cecum veiy large and showing through the 
foot (Plate 2, Fig. 5). Pedal and siphonal 
retractor muscles weak, their arrangement 
typical for the genus. 

Measurements. See Table 2. 

Remarks. On the basis of the shell and 
the mesoplax, this species is closely related 
to X concava Knudsen from the Gulf of 
Panama. It differs in being much smaller 



238 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



(none of the 19 specimens reached 4 mm 
in length, whereas Knudsen gives 8.6 mm 
for X concava) and in having a narrow me- 
soplax that does not extend above the um- 
bos. In addition, the excurrent and incur- 
rent siphons of X. gerda are the same 
length, are separate at the tip, and the ap- 
ertures of both siphons have four or five 
relatively large cirri. In X. concava, the si- 
phons are joined for their entire length 
and the excurrent siphon is slightly shorter. 
The type of chimney produced by X gerda 
is unlike any other known to date in this 
genus. 

Unfortunately, all the specimens of X 
gerda are small, extremely fragile, and 
rather poorly preserved. Consequently, it 
has been impossible to do anatomical work 
beyond that mentioned in the description. 
One specimen from which the valves were 
removed appeared to have an accessory 
genital organ similar to that described by 
Purchon (1941). 

Range. Probably throughout the Carib- 
bean in depths from about 283 to 2,072 m. 

Specimens Examined. BAHAMA ISLANDS; Ger- 
da, station 499, about 3 miles off Southwest Point, 
Great Bahama Island (26°37'N, 78°56'W) in 155 fath- 
oms (283 m). UNITED STATES, FLORIDA: Gerda, 
station 266, off Fowey Rocks, Florida Keys (25°39'N, 
79°58'W) in 185-187 fathoms (338-342 m). LESSER 
ANTILLES: Pillsbunj, station 944, 45 miles N of 
Port Louis, Guadeloupe Island (I6°32,2'N, 
61°36.8'W) in 364-421 m. PANAMA: Pillsbunj, sta- 
tion 328, about 25 miles N of Punta San Bias, Gulf 
of San Bias (9°55.8'N, 78°59.8'W) in 2,069-2,072 m. 

Xylophaga grevei Knudsen 
Plate 4 

Xylophaga grevei Knudsen, 1961, Galathea Report, 
5: 176, figs. 16-18 (Galathea, station 495, Banda 
Trench, south of Geram [5°26'S, 130°58'E] in 
7,250-7,290 m). Holotype, Zoological Museum, 
University of Copenhagen. 

Distinctive Characters. Mesoplax com- 
posed of two triangular plates that are flat 
dorsally, in contact the length of their iTie- 
dian edge, bent downward on their outer 
edge to meet the umbonal reflection, and 
lack a ventral portion. Posterior adductor 
muscle scar with oblique radiating impres- 



Table 3. Measurements of Xylophaga grevei. 



Length 

(mm) 



Height 

(mm) 



1,9 1.6 Galathea. station 444 

12.5 11.0 holotype 



sions. Siphons nearly the same length, the 
aperture of the excurrent siphon much 
smaller in diameter than the incurrent si- 
phon and with about 6 cirri; incurrent si- 
phon with about 35 small cirri. Young car- 
ried on the uinbonal area of the adult. 

Measurements. See Table 3. 

Remarks. Through the kindness of Jor- 
gen Knudsen, it was possible to borrow the 
preserved dredged wood from the Zoolog- 
ical Museum, University of Copenhagen. 
In a small piece taken by the Galathea at 
station 444, I found several additional 
speciiTiens of X. grevei. They are much i 
smaller than that figured by Knudsen, the 
anterior slope is much narrower, and the 
denticulated ridges are more widely 
spaced. However, they appear to be young 
but sexually mature specimens of that spe- 
cies. From one to five rather large young 
were attached posterior to the umbos on 
the dorsal surface of the parent shells. 

Xylophaga wolffi Knudsen, based on i 
only two specimens, also was fro in Gala- 
thea, station 444. It has valves, muscle 
scars, and siphons similar to those of X. 
grevei. The outstanding difference be- 
tween these species is the flat plates of the 
mesoplax of X. wolffi, which are set in 
tentlike fashion at an acute angle to each 
other. It has not been possible to compare 
the type of X. wolffi with this new material 
from Galathea, station 444, but it appears 
that these two forms (i.e., X. grevei and X. 
wolffi) may be equivalent to the condition 
found in X. clenchi Turner and Culliney, 
where occasional specimens have a bent, 
flat-topped mesoplax (Plate 4, Figs. 5, 6). 
Further collecting may show X. grevei and 
X. wolffi to be forms of a single species. 
The denticles on the ventral edge of the 
mesoplax of X. wolffi described by Knud- 



Xylophagainae • Turner 



239 



sen may be an age factor but more mate- 
rial is needed to prove this. 

Range. Mindanao Sea south to the Ban- 
da Trench, Banda Sea in depths from 
about 545 to 7,290 m. 

Specimens Examined. PHILIPPINE ISLANDS: 
Galathea, station 444, Sulu Sea, W of Basilan Island 
(7°54'N, 121°30'E) in 5,050 m. 

Xylophaga clench P Turner and Culliney 
Plates 5-8 

Xylophaga clenclii Turner and Culliney, 1971, Amer- 
ican Malacological Union Annual Report for 1970, 
p. 66 (U.S. NOO test site. Tongue of the Ocean, 
about 4 miles off northeastern tip of Andros Island, 
Bahama Islands [24°54'N, 77°49'W] in 1,737 m). 
Holotype, MCZ 316743; paratypes, MCZ 316744, 
316745. 

Distinctive Characters. Mesoplax small, 
composed of triangular, nearly flat plates 
lacking a basal portion; the two plates usu- 
ally meeting at an acute angle in frontal 
view. Burrow lined with a chimney of 
coarse, loosely consolidated fecal pellets. 
Young held on posterior dorsal surface of 
the adult. Excurrent siphon shorter than 
incurrent siphon and with two large papil- 
lae on either side. 

Description. Shell globose, reaching 14 
mm in length and 13.5 mm in height, thin, 
fragile, with a thin, light brown periostrac- 
um that is thickened along the dorsal and 
posterior margin of the valves. Umbos 
prominent and strongly incurved. Pedal 
gape angle about 108°. Beaked portion of 
the anterior slope sculptured with numer- 
ous denticulated ridges that are widely 
spaced in the young, becoming increasing- 
ly compacted toward the ventral margin in 
older specimens. Specimens 5 mm in 
length have up to 25 ridges. Posterior por- 
tion of the anterior slope rather narrow. 
Umbonal— ventral sulcus moderately to 
deeply impressed becoming shallower 
with age and bounded posteriorly by a 
broad, low, rounded ridge. Disc and pos- 



^ Named for Dr. William J. Clench, Curator of 
Mollusks, 1926—1966, Museum of Comparative Zo- 
ology, Harvard University. 



terior slope sculptured with uniform, rath- 
er pronounced growth ridges that are par- 
ticularly prominent in young specimens. 

Inner surface of valves smooth and glis- 
tening. Umbonal-ventral ridge prominent, 
distinctly segmented, and with a large ven- 
tral condyle. Chondrophore and internal 
ligament well developed. Posterior adduc- 
tor muscle scar kidney-shaped in outline 
and with irregular transverse, often anas- 
tomosing impressions that become more 
numerous with age. Pedal retractor scar 
more or less oval in outline and located 
just anterior to the embayment in the pos- 
terior adductor scar. Siphonal retractor 
scars small, not impressed, located just 
posterior to the umbonal-ventral ridge 
about midway between the umbo and the 
ventral condyle. 

Mesoplax small, composed of two flat to 
slightly arched triangular plates lacking a 
ventral portion, set in a tentlike fashion 
and held in place by the periostracum and 
the anterior adductor muscle. Plates usu- 
ally meet dorsally at 90° (Plate 5, Fig. 3), 
occasionally at an obtuse angle. They are 
rarely bent longitudinally (Plate 8, Fig. 3). 
Outer surface of plates sculptured with 
ridges paralleling the anterior margin and 
lining up with the denticulated ridges of 
the valves w^here they come in contact. 

Siphons short, extending only about one 
third the length of the valves. Excurrent 
siphon about one half the diameter of and 
slightly shorter than the incurrent siphon, 
with a well-developed sphincter muscle 
surrounding the aperture and two large 
papillae on each side (Plate 7, Fig. 3). In- 
current siphon has minute cirri surround- 
ing the aperture and a second set of larger 
cirri anterior to them within the siphon 
(Plate 6, Fig. 4). 

Posterior end of burrow lined with 
coarse, loosely compacted fecal pellets. 
Young held by byssus threads on the pos- 
terior dorsal surface of the adult shell 
(Plate 7, Figs. 1, 2). 

Measurements. See Table 4. 

Remarks. This species is most closely re- 
lated to X africana Knudsen from the 



240 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 4. Measurements of Xylophaga clenchi. 



Length 


Height 




(mm) 


(mm) 


Locatioii 


2.3 


2.1 


Ingolf, Station 67 


3.2 


2.0 


Atlantis II, station 124 


3.5 


3.5 


holotype 


4.0 


3.5 


Tongue of the Ocean 


5.0 


4.8 


Tongue of the Ocean 


10.0 


9.5 


Tongue of the Ocean 


10.0 


10.0 


Pillsbury, station 104 


14.0 


13.5 


Pillshury, station 394 



Gulf of Guinea, West Africa (1°42'N, 
7°51'E in 2,550 m), but the mesoplax dif- 
fers in having a longer median line where 
the two plates meet, a weaker sculpture, 
and in lacking the rounded nodule on the 
ventral surface. Unfortunately, the valves 
of the two known specimens of X. africana 
were too fragixientaiy to allow comparisons 
on this basis. The siphons of these two 
species are similar but X clenchi differs in 
having only two rather than three large cir- 
ri on either side of the excurrent siphon 
and in having two rings of small cirri bor- 
dering the aperture of the incurrent si- 
phon. The incurrent siphon of X. africana 
lacks cirri, according to Knudsen (1961). 
Two other species, X wolffi Knudsen from 
the Sulu Sea and X murrayi Knudsen 
from off Zanzibar, also have similar dorsal 
plates. The former differs from X. clenchi 
in having denticles on the basal margin of 
the mesoplax with corresponding denticles 
on the umbonal reflection. In addition, on 
each side of the excurrent siphon of X. 
wolffi there are "7 finger-like tentacles on 
a common base and somewhat larger ten- 
tacle dorsally." The siphons of X. murrayi 
differ in having a circle of about 35 "short 
tentacles" surrounding both openings. 
Knudsen (1961) compared X. wolffi with 
X. supplicata Taki and Habe but exami- 
nation of paratypes of X. supplicata re- 
ceived from Habe showed that species to 
have tubules on the mesoplax. (See also 
under X. supplicata Taki and Habe.) 

Xylophaga clenchi Turner and Culliney 
also appears to be closely related to X. 
panamensis Knudsen but the latter differs 



in having the plates of the mesoplax 
sinooth. The posterior adductor muscle 
scar of X. panamensis is oval and irregu- 
larly lobed anteriorly, the weak umbonal- 
ventral ridge is bounded posteriorly by a 
groove, and the condyles are lacking. The 
siphons of X. panam,ensis are unknown. \ 

The variation in the inesoplax of X. clen- 
chi is interesting and presents some prob- 
lems. In most specimens, the plates are flat 
or nearly so and none has a ventral portion 
or cavity of any kind. The angle at w^hich 
the plates meet dorsally is typically acute; 
however, specimens from shallower waters 
(i.e., less than 900 m) occasionally have 
plates that meet at an obtuse angle. Two 
of the five specimens from Atlantis II, sta- 
tion 124, have the dorsal plate bent lon- 
gitudinally at a right angle (Plate 8, Fig. 
3), whereas others are typical. Because the 
shell characteristics of these specimens fit 
within the range of variation of the typical 
forms, they are considered to be ecologic 
variants. Unfortunately, none of the spec- 
imens from shallower water has extended 
siphons, so comparisons cannot be made 
on that basis. 

Although the holotype of X. clenchi is 
small, it was selected because it was the 
most nearly perfect and the only specimen 
for which dorsal plates, young, siphons, 
and chimney all were present. The large 
specimen from IW Pillsbury, station 394, 
and the two from Pillsbury, station 104, 
were dead when collected. At least one 
specimen from all other localities was alive 
at the time of collection. The specimens 
froiTi the Tongue of the Ocean were from 
test panels submerged by the NOO from 
4 April 1962 to 17 February 1965. For a 
description of the test site see DePalma 
(1969). One specimen taken by the IW 
Atlantis II at station 119 and three speci- 
mens taken at station 131 were from epi- 
benthic sled hauls in which no wood or 
plant material was found. The specimens 
were alive and apparently normal although 
they were badly crushed, for they were mi- 
nute and veiy delicate. This raises the 
question as to whether or not at least some 



Xylophagainae • Turner 



241 



species of Xylophaga can survive in a firm 
muddy bottom if wood is not available. So 
far as known, these are the only living 
specimens of Xylophaga not taken from 
wood or other plant material, except those 
that were boring in plastic at the navy test 
site off California and possibly those spec- 
imens of Xylophaga foliata Knudsen that 
came from a station in Macassar Strait 
from which it was reported that no wood 
was taken. The specimens from off Iceland 
were removed from wood dredged by the 
Ingolf Expedition in 1896. All specimens 
were small, the largest being 2.5 mm in 
length. Many had from 1 to 10 large young 
attached to the valves in the umbonal area 
(Plate 7, Fig. 2). This record extends the 
range of the species far to the north but it 
has been impossible to find any characters 
to distinguish these specimens from those 
occurring to the south. 

The difference in the size of the young 
attached to the dorsal surface of the spec- 
imens figured in Plate 5, Figures 1 and 2, 
and Plate 7, Figures 1 and 2, reflects the 
age of the lan^ae, with those in the latter 
plate having well-developed umbos and 
appearing fully mature. It is interesting to 
note the position of the larvae on the two 
adults and to speculate that perhaps the 
larvae move gradually toward the umbos 
as they mature. 

Range. From off Iceland south to Ve- 
nezuela in depths ranging from 35 to 
4,862 m. 

Specimens Examined. ICELAND: Ingolf, station 
67, S of Eyrabakki (61°30'N, 22°30'W) in 1,836 m. 
UNITED STATES, NEW JERSEY: Albatross, station 
2550, about 160 miles E of Barnegat Bay (39°44'N, 
70°30'W) in 1,977 m. VIRGINIA: Albatross, station 
2731, off Cape Henry (36°45'N, 74°28'W) in 1,428 
m. NORTH CAROLINA: Atlantis II, station 131, 
about 420 miles E of Currituck Sound (36°28.9'N, 
67°58.2'W) in 2,178 ni; off Cape Hatteras (35°44'N, 
75°15'W) in 35 m. GEORGIA: Pillsburij, station 104, 
about 80 miles SE of Brunswdck (3r00'N, 79°50'W) 
in 247 m. BAHAMA ISLANDS: Tongue of the 
Ocean, about 4 miles off NE tip of Andros Island 
(24°54'N, 77°49'W) in 1,737 m; Tongue of die Ocean 
(24°53.2'N, 77°40.2'W) in 2,066 m. BERMUDA: At- 
lantis II, station 124, about 750 miles E of Cape 
Charles (due N of Bermuda) (37°26'N, 63°59'W) in 



Table 5. Measurements of 
Xylophaga supplicata. 



Length Height 

(mm) (mm) 



8.5 



8.8 



parat>pe MCZ 194820 



4,862 m; Atlantis II, station 119, just S of Bermuda 
(32°15'N, 64°32'W) in 2,095-2,223 m. PANAMA: 
Pillsbunj, station 328, about 25 miles off Punta San 
Bias, Gulfo San Bias (9°55.8'N, 78°59.8'W) in 420- 
640 m. VENEZUELA: Pillsbunj, station 719, about 
100 miles N of Pertigalete Bay (11°35'N, 64°35.4'W) 
in 770-890 m. 

Xylophaga supplicata Taki and Habe 
Plate 9 

Metaxi/lophaga supplicata Taki and Habe, 1950, Il- 
lustrated Catalogue of Japanese Shells No. 7, p. 47, 
text-figs. 1, 2 (Tosa Bay, Shikoku, Japan, in 100 
fathoms). Holotype, T Habe Collection; paratype, 
MCZ 194820; Knudsen 1961, Galathea Report, 5: 
188. 

Distinctive Characters. Mesoplax com- 
posed of two flat triangular plates set at a 
sharp angle to each other, dorsal margin 
more than one half the total length, and 
with minute tubules at the posterior end. 
The tubules sit deep within the cavity 
formed by the umbos and cannot be seen 
when the shell is viewed anteriorly. Chon- 
drophore of left valve with a large tooth 
(Plate 9, Figs. 5, 6). 

Measurements. See Table 5. 

Remarks. Taki and Habe (1950) de- 
scribed the dorsal plate of X supplicata 
simply as a "small triangular protoplax 
[ = mesoplax]" that is attached vertically. A 
paratype received from Dr. Habe has small 
tubes on the dorsal posterior end of the 
mesoj^lax. 

Xylophaga bniuni Knudsen, 1961, from 
the Mindanao Sea is very close to if not 
synonymous with X supplicata. Unfortu- 
nately, Taki and Habe did not mention the 
tubes in their original description and 
Knudsen did not see the types. Also, un- 
fortunately, Knudsen had only a single 
specimen; hence, it will be necessaiy to 
obtain more material before a definite 
statement can be made concerning the sta- 



242 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



tus of X bniuni. (See also Remarks under 
Xylophaga ivhoi Turner n. sp.) 

Specimens Examined. JAPAN: Tosa Bay, Shikoku, 
in 100 fathoms (183 m). " 

Xylophaga whoh Turner new species 
Plates 10, 11 

Holotijpe. MCZ 275015. 

Type Locality. R/V Atlantis, station 
3471, off Cardenas, Matanzas Province, 
Cuba (23°21'N, 80°56'W) in 500 fathoms 
(914 m). 

Distinctive Characters. Mesoplax com- 
posed of two flat triangular plates with a 
short median line, set at a shai-p angle to 
each other and with a large hollow tube 
extending outward from the posterior dor- 
sal surface of each plate. 

Description. Shell globose, reaching 7 
mm in length and 6.2 mm in height, thin, 
fragile, and with a veiy thin, light tan per- 
iostracum. Pedal gape angle about 95°. 
Beaked portion of the anterior slope sculp- 
tured with numerous low denticulated 
ridges; the posterior portion with fine in- 
distinct ridges. Umbonal-ventral sulcus 
only slightly impressed and irregularly 
sculptured. Sculpture on the disc and pos- 
terior slope consisting of pronounced 
growth ridges. Umbonal reflection narrow, 
closely impressed near the umbos, free 
and vertical anteriorly. 

Inner surface of valves smooth and shin- 
ing. Umbonal-ventral ridge becoming ev- 
ident only near the ventral margin but rap- 
idly increasing in size to a large ventral 
condyle. Chondrophore and internal liga- 
ment well developed; chondrophore of the 
left valve with a small tooth on the poste- 
rior upper margin. Posterior adductor 
muscle scar large, elongate, oval in outline, 
irregularly sculptured, set high on the pos- 
terior slope and well in from the posterior 
margin. Pedal retractor scar rather large, 
elongate, somewhat irregular, and located 



Table 6. Measurements of Xylophaga whoi. 



Leiiytli 

(mil.) 


Height 
(mm) 


Location 


2.5 


2.3 


Gerda, station 266 


3.1 


3.0 


Gerda, station 266 


6.9 
7.0 


6.2 
6.8 


holotype 

Pillsbiii-y, station 394 


7.2 


7.0 


Pillsbury, station 944 


9.5 
12.5 
13.8 
14.9 


9.0 
11.5 
13.1 
14.0 


Pillsbury, station 944 
Pillsbury, station 944 
Pillsbury, station 944 
Pillsbury, station 944 



just anterior to the posterior adductor scar. 
Siphonal retractor scar not impressed. 

Mesoplax composed of two triangular 
plates that are set at an acute angle to each 
other dorsally and have two large, hollow, 
tubular projections extending from the 
posterior dorsal margin (Plate 10, Figs. 3— 
10). Length of dorsal margin less than one 
half the length of the plate, outer surface 
sculptured with distinct ridges paralleling 
the anterior margin; inner surface smooth, 
with a small internal opening into the tube 
(Plate 10, Figs. 11, 12). 

Siphons short, of about equal length, 
and probably not capable of extending 
more than one half the length of the shell. 
Aperture of incurrent siphon large and ap- 
parently lacking cirri, that of the excurrent 
siphon small and with a few fine cirri on 
each side ventrally. 

Measiirenients. See Table 6. 

Remarks. Knudsen (1961) described 
three species of Xylophaga with tubulate 
mesoplaxes, and studies for this report 
have shown that X supplicata Taki and 
Habe also is tubulate. Three of these spe- 
cies are from the western Pacific (X. tuh- 
ulata Knudsen from Macassar Strait, X. 
briiiini Knudsen from the Mindanao Sea, 
and X. supplicata Taki and Habe from Ja- 
pan), and one from the eastern Pacific (X. 
obtusata Knudsen from the Gulf of Pana- 
Xijlophaga whoi Turner n. sp. is the 



ma) 



' An acronym for Woods Hole Oceanographic In- 
stitution, whose research vessel Atlantis collected the 
holotype. 



first tubulate species reported from the At- 
lantic. It is most closely related to X. bni- 
uni Knudsen from the Mindanao Sea but 
differs in that the mesoplax has a propor- 



Xylophagainae • Turner 



243 



tionately shorter dorsal margin and larger 
tubes. In addition, the posterior adductor 
muscle scar is set well in from the poste- 
rior margin and the umbonal-ventral ridge 
is evident only near the ventral margin of 
tlie shell but increases rapidly to a large 
condyle. Xtjlophaga whoi Turner n. sp. dif- 
fers from X. obtusata Knudsen in the po- 
sition of the posterior adductor scar, the 
shape of the mesoplax, the larger tubes on 
the mesoplax, the narrower anterior slope, 
and smaller umbonal reflection. The third 
tubulate species described by Knudsen, X. 
tubulata froni Macassar Strait, differs from 
X. whoi in having very large tubules that 
extend to the anterior margin of the me- 
soplax and in having the plates bent lon- 
gitudinally at a right angle so they are flat 
dorsally rather than meeting at an acute 
angle. Examination of a paratype specimen 
of X. supplicata Taki and Habe, 1950, from 
Japan shows that this species has minute 
tubes on the mesoplax similiar to those in 
X. briiiini and it may be synonymous with 
that species. (See also under X supplicata 
Taki and Habe.) 

The specimen of X. whoi Turner n. sp. 
taken by the RA^ Pillsbiinj from off Punta 
Piedras, Colombia, had 13 young attached 
to the umbonal area (Plate 11, Fig. 2). 
Among the other species in this group 
(Group 3), Knudsen (1961) reported that 
the type of X obtusata had two young and 
the type of X. bruuni had four young on 
the dorsal surface of the adult shell, 
whereas X. tubulata had five young at the 
ventral base of the siphons in a depression 
of the mantle tissue. The young at the base 
of the siphons of X. tubulata were very 
small, about one half the size of those on 
X. bixiuni. They possibly had only recently 
been extruded from the excurrent siphon 
of the adult and had not yet crawled to the 
dorsal surface of the adult. It has not been 
proven that the juveniles clinging to the 
mantle and shells of adult Xylophaga are 
definitely the young of the specimen to 
which they are attached, and this has been 
questioned by some workers. However, 
until the species concerned are cultured in 



the laboratory, it is the only assumption 
that can be made safely. The only way the 
young of another specimen or species 
could get into the burrow would be via the 
incurrent siphon. For the veliger lai'vae of 
another specimen to get to the umbonal 
area, they either would have to pass 
through the gills into the epibranchial 
chamber and then out the excurrent si- 
phon, or go through the digestive tract to 
the excurrent siphon. Neither of these al- 
ternatives seems likely. It is, of course, pos- 
sible that the young of X. tubulata com- 
plete their development at the base of the 
siphons of the parent. 

Range. From off southern Florida to 
Colombia in depths from about 336 to 
910 m. 

Specimens Examined. UNITED STATES, ELOR- 
IDA; Gerda, station 266, off Eowey Rocks (25°39'N, 
79°58'W) in 185-187 fathoms (338-345 m). CUBA: 
Atlantis, station 3471, off Cardenas, Matanzas Prov- 
ince (23°21'N, 80°56'W) in 500 fathoms (914 m). 
LESSER ANTILLES: PiUsbunj, station 944, 45 miles 
N of Port Louis, Guadeloupe Island (16°32.2'N, 
61°36.8'W) in 364-421 m. COLOMBIA: PiUsbunj, 
station 394, off Punta Piedras (9°28'N, 76°26'W) in 
230-350 fathoms (419-640 m). 

Xylophaga profunda Turner new species 
Plates 12, 13 

Holotijpe. MCZ 316751. 

Type Locality. Tongue of the Ocean, off 
NE tip of Andros Island, Bahama Islands 
(25°54'N, 77°49'W) in 1,722 m. From test 
panel submerged from 26 July 1962 to 17 
February 1965. 

Distinctive Characters. Valves with a 
well-impressed umbonal-ventral sulcus, 
bounded posteriorly by a low broadly 
rounded ridge. Posterior slope sculptured 
with narrow, concentric grooves. Mesoplax 
of two triangular plates, lacking a basal 
portion, set at an acute angle to each other, 
folded longitudinally with the anterior 
margin bent inward and with a small pore 
in each anteriorly. Umbonal-ventral ridge 
on inner surface of valves narrow, high, 
and strongly segiTiented. 

Description. Shell globose, reaching 
about 11 mm in length, thin, fragile, and 



244 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 7. Measurements of 
Xylophaga profunda. 



Lengtli Height 
(mm) (mm) 



5.9 5.6 holotype 

Tongue of the Ocean, off NE tip of 
8.0 7.9 Andros Island, Bahama Islands 

Tongue of the Ocean, off NE tip of 
9.5 8.8 Andros Island, Bahama Islands 

Tongue of the Ocean, off NE tip of 
10.0 9.8 Andros Island, Bahama Islands 

Tongue of the Ocean, off NE tip of 
10.9 10.8 Andros Island, Bahama Islands 



with a relatively heavy, light brown perios- 
tracum. Pedal gape angle about 95°. 
Beaked portion of anterior slope sculp- 
tured with numerous strong, denticulated 
ridges that are more closely spaced toward 
the ventral margin in adults. Umbonal— 
ventral sulcus narrow, deep, bounded pos- 
teriorly by a low, broad, rounded ridge. 
Posterior slope sculptured with widely 
spaced, incised grooves. Umbonal i^eflec- 
tion narrow, high, and strongly segmented. 
Incised grooves of posterior slope ex- 
pressed internally as faint ridges just pos- 
terior to the umbonal— ventral ridge gi\^ng 
a "back bone and ribs effect." Posterior ad- 
ductor muscle scar broadly oval, set high 
on the posterior slope, and lightly marked 
with irregularly anastomosing depressions. 
Pedal retractor scar lightly impressed, 
dumbell-shaped, and located just anterior 
to the midportion of the posterior adduc- 
tor. Siphonal retractor scar large, elongate, 
lightly impressed, and located about mid- 
way between the posterior adductor and 
the umbonal— ventral ridge. Chondrophore 
large, that of the left valve with two small 
teeth on the posterior edge. 

Mesoplax composed of two triangular 
plates lacking a basal portion, set at an 
acute angle to each other, folded longitu- 
dinally with the anterior margin bent in- 
ward and with a small pore in each ante- 
riorly at the outer edge of the fold. 

Measurements. See Table 7. 

Remarks. This species is superficially 
similar to X abijssonini Dall but differs in 



having a low, broad, rounded ridge poste- 
rior to the umbonal— ventral sulcus, in lack- 
ing the deep groove posterior to the um- 
bonal— ventral ridge on the inner surface, 
in having a pronounced sculpture on the 
posterior slope, and in being larger. In ad- 
dition, the inesoplax of X. profunda lacks 
a ventral portion, has only a single pore in 
each plate, does not produce tubes, and 
appears narrow when viewed anteriorly. 

Xylophaga profunda Turner n. sp. is 
probably most closely related to X lohata 
Knudsen from the Sulu Sea. Xijlophaga 
profunda differs in having a mesoplax that 
is much longer than wide, whereas in X 
lobata the structure is wider than long. 
Both species have a strongly segmented 
umbonal— ventral ridge and a similarly 
marked posterior adductor scar, although 
in X. profunda the markings are more ex- 
tensive and elaborate. Knudsen (1961) 
compared his X. galatheae from the Tas- 
man Sea with X lobata, noting differences 
in the mesoplax and muscle scars. Unfor- 
tunately, he had only a single complete 
specimen of X. galatheae, so he could not 
determine range of variation. Xylophaga 
galatheae possibly is a young X. lobata but 
more material from inteivening areas is 
needed to ascertain this. 

On the basis of the general shape and 
attachment of the dorsal plates, X. profun- 
da, along with X. lobata and X. galatheae, 
appear to belong in species Group 3, al- 
though X. profunda is not closely related 
to any species in the group and is placed 
here tentatively. The major differences in- 
clude the sculpturing of the posterior ad- 
ductor muscle scar, the presence of ante- 
rior pores, and the lack of posterior tubes 
on the mesoplax. The chondrophore of the 
left valve of X. profunda has two small 
teeth, a character that relates it to X. ivhoi 
and X. supplicata, both of which have a 
single large tooth. Unfortunately, Knudsen 
did not mention the presence of teeth in 
the three tubulate species he described. 
Consequently, it cannot be stated definite- 
ly that this is characteristic of the group. 

The siphons of X. profunda are short. 



Xylophagainae • Turner 245 



probably of the same length and capable 
of complete retraction within the valves. 
Unfortunately, none of the specimens was 
sufficiently well preserved for anatomical 
work. Four specimens carried young on 
the umbonal area, the smallest had 5 and 
the largest had 75. The young have pro- 
nounced umbos, distinct concentric sculp- 
ture, and measure about 0.30 mm in 
length. 

Xylophaga profunda Turner n. sp. is 
known from five specimens taken from 
one panel submerged in the Tongue of the 
Ocean at 25°54'N, 77°49'W, and from 14 
specimens taken from an asbestos-backed 
panel submerged at 24°53.2'N, 77°40.2'W. 

Specimens Examined. BAHAMA ISLANDS: 
Tongue of the Ocean, off NE tip of Andros Island 
(25°54'N, 77°49'W) in 1,722 ni; Tongue of the Ocean, 
Tower 3 (24°53.2'N, 77°40.2'W) in 2,066 m. 

Xylophaga abyssorum Dall 
Plates 14-16 

Xylophaga abyssorum Dall, 1886, Bulletin Museum 
of Comparative Zoology, 12: 317, pi. 9, fig. 7, 7a 
{Blake, station 215, off St. Lucia, Lesser Antilles 
[13°51'N, 61°03'W] in 226 fathoms). Holotype, 
MCZ 8135; Turner, 1955, Johnsonia, 3(34): 156, pi. 
93. 

Distinctive Characters. Valves with a 
proiTiinent ridge just posterior to the um- 
bonal— ventral sulcus and with a slightly to 
strongly concave profile posterior to the 
ridge when viewed dorsally Umbonal— 
ventral ridge on the inner surface devel- 
oped only near the ventral condyle and 
bounded posteriorly by a deep groove. 
Mesoplax composed of two more or less 
triangular plates having a ventral portion 
and a variously lobed dorsal portion; lobes 
varying with age and coalescing to form 
pores or tubes (Plate 15, Figs. 4, 5). 

Description. Shell globose but appear- 
ing constricted posteriorly when viewed 
dorsally, reaching 5.5 mm in length, thin, 
fragile, and with a thin, light brown per- 
iostracum. Pedal gape angle about 95°. 
Beaked portion of the anterior slope sculp- 
tured with fine, denticulated ridges, widely 
spaced in young specimens but compacted 



toward the ventral margin in older speci- 
mens. Umbonal— ventral sulcus narrow, 
rather shallow, and with a narrow promi- 
nent ridge just posterior to it. Profile of 
valves concave posterior to the ridge when 
viewed dorsally. Posterior slope sculptured 
with faint concentric growth lines, umbo- 
nal reflection erect, the ventral margin of 
the mesoplax meeting it and attached by a 
periostracal fold. Umbos inflated, particu- 
larly in young specimens. 

Inner surface of valves white and glazed. 
Muscle scars well marked. Posterior ad- 
ductor scars somewhat pear-shaped, set 
well in from the posterior margin, and 
marked with irregular, elongate depres- 
sions extending inward from the posterior 
margin. Anterior adductor scar covering 
most of the umbonal reflection. Pedal re- 
tractor scar nearly circular and located just 
anterior to the widest position of the pos- 
terior adductor scar. Siphonal retractor 
scar small, elongate, lightly impressed, and 
located anterior and ventral to the pedal 
retractor. Umbonal— ventral ridge not well 
developed except near the ventral condyle 
but bounded posteriorly by a deep groove 
that is a reflection of the external ridge. 
Chondrophore well developed, internal 
ligament strong. 

Mesoplax variable, with a well-devel- 
oped, more or less triangular basal portion, 
occasionally with lateral notches in young 
speciniens. Dorsal portion developing and 
vaiying with age; in young specimens, con- 
sisting of lobes extending anteriorly from 
the posterior ends of the plates (Plate 14, 
Fig. 2) followed by lobed lateral folds 
(Plate 14, Figs. 3-5, 7), which grow and 
eventually coalesce to form pores or short 
tubes (Plate 15, Figs. 4, 5, and Plate 16, 
Figs. 4, 5). Aperture of the tubes in living 
specimens covered by a periostracal mem- 
brane. 

Siphons short, about equal in length, 
and capable of complete retraction within 
the valves. Diameter of the incurrent si- 
phonal aperture about twice that of the ex- 
current siphon. The margins of both ap- 
pear to have cirri. 



246 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 8. Measurements of 
Xylophaga abyssorum. 



Length Height 
(mm) (mm) 



3.0 2.6 Pillsbury, station 944 

3.0 2.9 Tongue of the Ocean, off NE tip of 
Andros Island, Bahama Islands 

3.3 3.0 Pillsbunj, station 944 
3.9 3.2 Pillsbunj, station 944 
4.0 3.5 holotype 

4.0 3.6 Gerda, station 266 

5.4 4.5 PiUsbun/, station 944 



Measurements. See Table 8. 

Retnarks. Dall based his description of 
X abyssorum on two isolated valves of a 
dead specimen, with the main distinguish- 
ing character being the pronounced ridge 
posterior to the umbonal— ventral sulcus. 
All specimens that have been assigned to 
this species since that time have been so 
named on the basis of this ridge. Among 
the specimens taken from boards sub- 
merged in the Tongue of the Ocean off 
Andros Island, Bahama Islands, by the 
U.S. NOO (DePalma, 1969), was a single 
specimen the valves of which coincided 
with those of the holotype. Two specimens 
were taken from wood dredged off Gua- 
deloupe Island, Lesser Antilles. All of 
these specimens have a pronounced ridge, 
concave posterior slope, and lobed meso- 
plax. With this new material, it has been 
possible to redescribe X abyssonnn giving 
the characters of the mesoplax and a range 
of variation. Plate 14, Figure 6, illustrates 
the dorsal view of the right valve of the 
holotype of X. abyssorum; Figure 3 illus- 
trates the dorsal view of the right valve of 
the specimen from the Tongue of the 
Ocean. These two valves so closely resem- 
ble each other that, except for a slight dif- 
ference in size, they could be from the 
same specimen. Variations in the valves 
and dorsal plates of specimens from Flor- 
ida and Guadeloupe Island are illustrated 
on Plates 14-16. Plate 15, Figure 5, illus- 
trates the position in which the young are 
carried on the adult. The largest number 
of young on any one specimen was five; 



the average measurement of the young 
was 0.3 mm in length. 

Xylophaga abijssorum has a small shell 
but is a distinctive species, particularly in 
the adult stage. It is probably most closely 
related to X. lobata Knudsen from the 
Sulu Sea but differs in having a sharp ridge 
posterior to the umbonal— ventral sulcus 
and a far more elaborately lobed mesoplax 
in the adult. From X. profunda Turner n. 
sp., the only other species with lobes on 
the dorsal portion of the mesoplax, X 
abyssorum differs in having broad dorsal 
plates with two or more pores or tubes in 
each. In addition, X. profunda is a much 
larger species, the umbonal— ventral ridge 
on the outer surface of the valves is broad 
and rounded, and the deep groove on the 
inner surface is lacking. The posterior ad- 
ductor muscle scars of X. profunda are 
lightly and irregularly marked, whereas the 
inuscle scars of X. abyssorum are well 
marked. In the young stage, the ixiesoplax 
of X abyssorum is similar to that of X. baij- 
eri Turner n. sp. and X. profunda, but the 
valves of these species do not have a pro- 
nounced ridge. The shape of the valves of 
X. abyssorum somewhat resembles those 
of X. japonica Taki and Habe but the me- 
soplax of that species is ear-shaped and not 
lobed. (See under X. japonica.) 

Range. Based on the valves of dead 
specimens lacking a mesoplax, the range of 
this species extends from off Atlantic City, 
New Jersey, south to St. Lucia, Lesser An- 
tilles (Turner, 1955: 157). Living speci- 
mens are known only from Florida, the 
Bahamas, and Guadeloupe Island, Lesser 
Antilles, in depths ranging from 342 to 
1,722 m. 



Specimens Examined. UNITED STATES, FLOR- 
IDA: Gerda, station 266, off Fowey Rocks (25°38'N, 
79°58'W) in 185-187 fathoms (338.3-342 m). BA- 
HAMA ISLANDS: Tongue of the Ocean, about 4 
miles off NE tip of Andros Island (24°54'N, 77°47'W) 
in 1,722 m. LESSER ANTILLES: Pillsbunj, station 
944, 4.5 miles N of Port Louis, Guadeloupe Island 
(16°32.2'N, 61°36.8'W) in 360-420 m. 



XylOPHAGAINAE • Turner 



247 



Table 9. Measurements of 
Xylophaga duplicata. 



Length 
(mm) 



Height 
(mm) 



3.7 3.5 holoty].ie 

5.2 5.0 Gulf of Tehuantepec, Me.\ico 

5.5 5.0 Gulf of Tehuantepec, Mexico 



Xylophaga duplicata Knudsen 
Plate 17 

Xylophaga duplicata Knudsen, 1961, Galathea Re- 
port, 5: 175, figs. 14, 15 {Galathea, station 745, 
Gulf of Panama [7°15'N, 79°25'W] in 915 m). Ho- 
lotype. Zoological Museum, University of Copen- 
hagen. 

Distinctive Characters. Plates of the 
mesoplax elongate oval, with a large ven- 
tral portion, somewhat inflated, diverging 
anteriorly, and standing off from the sur- 
face of the valves. Anterior adductor mus- 
cle extending into the cavity of the meso- 
plax. Anterior slope scLilptured with nu- 
merous exceedingly fine, closely set den- 
ticulated ridges. Umbonal reflection 
narrow and erect. Umbonal— ventral sulcus 
narrow, slightly impressed, and bounded 
posteriorly by a faint, rounded ridge. Pos- 
terior adductor muscle scar oval and 
smooth. Umbonal— ventral ridge narrow, 
high, and segmented; ventral condyle 
small. Siphons of equal length, united ex- 
cept at the tip, the posterior three-fourths 
covered with a brown periostracal sheath; 
siphonal openings (apertures set on two 
short tubes) each with six to eight cirri. 

Measurements. See Table 9. 

Remarks. At the time Knudsen de- 
scribed this species he had only two spec- 
imens from the Gulf of Panama. Two ad- 
ditional specimens received from D. Shas- 
ky were taken by the San Juan Expedition 
at station N-12 from a sunken log dredged 
in 60 fathoms (109 m) in the Grflf of Te- 
huantepec, Mexico (15°08'N, 93°28'W). 
Although the specimens were in rather 
poor condition, the valves and siphons 
agree with those described by Knudsen. 
The mesoplax stands off from the surface 
of the shell and, as stated by Knudsen, is 



double, that is, has a basal portion, and the 
anterior muscle extends into the cavity of 
the mesoplax. The siphons are the same 
length, combined in a common sheath ex- 
cept at the tip, and are covered with a thin, 
brown periostracum. The white spots 
mentioned by Knudsen are not evident, 
but the few cirri surrounding the siphonal 
openings are similar. 

Range. This record extends the geo- 
graphic range of the species about 500 
miles to the north. The depth range is 
from 109 to 915 m. 

Xylophaga muraokai^ Turner new species 
Plates 18, 19 

Holotype. MCZ 316746; paratypes, 
MCZ 316747, 316748, 316749, 316750. 

Type Locality. U.S. Naval Civil Engi- 
neering Laboratory Test Site I, about 81 
miles SW of Port Hueneme, California, 
or about 25 miles S of San Miguel Is- 
land, Santa Barbara Islands (33°44'N, 
120°45'W) in 5,640 ft (1,720 m). The ho- 
lotype was taken from a panel exposed on 
STU 1-2 from October 1963 to October 
1965. 

Distinctive Characters. Plates of the 
mesoplax wedge-shaped, the basal portion 
large and the small dorsal portion covering 
only the posterior part of the muscle. Si- 
phons smooth, of rmequal length, the ex- 
current slightly shorter and with 8-10 
proininent cirri. 

Description. Shell globose, valves reach- 
ing 14 mm in length and 13 mm in height; 
the width of apposed valves about 12 mm. 
Valves thin but strong, white, with a thin 
clear, transparent to bright yellow perios- 
tracum. Angle of the pedal gape about 
115°. Anterior slope sctilptured with nar- 
row, finely denticulated ridges that are 
widely spaced during early growth, becom- 
ing more closely spaced toward the ventral 
margin in older specimens. Holotype with 
17 ridges on the anterior slope. Umbonal— 
ventral sulcus rather narrow, only slightly 



' Named for James Muraoka, Biologist, U.S. Naval 
Engineering Laboratory, Port Hueneme, California. 



248 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



depressed and sculptured widi fine growth 
lines. Disc and posterior slope sculptured 
with growth lines only. 

Inner surface of valves smooth and shin- 
ing. Unibonal— ventral ridge narrow and 
marked with distinct transverse ridges that 
appear knobby in some specimens. Chon- 
drophore and internal ligament well de- 
veloped. Posterior adductor muscle scar 
large, elliptical, covering most of the pos- 
terior slope, and marked with irregular, 
faint depressions that generally cross the 
short axis of the scar. Pedal retractor mus- 
cle scar broadly oval in outline and located 
just anterior to and about midway on the 
posterior adductor scar. In some speci- 
mens, the two scars appear to be adjacent. 
Siphonal retractor muscle scars not clearly 
defined but located just posterior to the 
umbonal— ventral ridge about midway dor- 
soventrally. Ventral adductor muscle scar 
long, narrow, and traversing the ventral 
end of the umbonal— ventral ridge. Ante- 
rior adductor muscle scar covering the 
umbonal reflection and the ventral flange 
of the mesoplax. 

Mesoplax small and divided. The ventral 
portion is a large, cuived, wedge-shaped 
shield, fitting closely against the surface of 
the valves. Posteriorly, the mesoplax curves 
upw^ard and foi"Avard covering the posterior 
portion of the anterior adductor muscle 
(Plate 18, Figs. 3-6). 

Siphons smooth, united, and capable of 
extending over three times the length of 
the valves. Excurrent siphon only slightly 
shorter than the incurrent siphon. Aper- 
ture of the incurrent siphon bordered by 
18—22 long, slender cirri that, in preserved 
specimens, curl inward and are not easily 
seen. At the base of these are numerous 
small cirri and, anterior to them on the 
inner wall of the siphon, 8—10 broad, 
branched cirri. Distal margin of excurrent 
siphon with two broad, short cirri on each 
side adjacent to the incurrent siphon, and 
8—10 long, slender cirri, the longest locat- 
ed middorsally, with those on either side 
becoming progressively shorter. 

The anterior adductor muscle inserts 



Table 10. Measurements of 

XYLOPHAGA MURAOKAl. 



Length 


Height 




(mm) 


(mm) 


Location 


6.5 


6.2 


paratype 


8.5 


8.0 


paratype 


11.8 


12.0 


paratype 


12.0 


11.5 


paratype 


12.0 


11.9 


paratype 


12.8 


12.8 


holotype 


14.0 


13.0 


paratype 



both on the anterior reflection and on the i 
basal portion of the mesoplax. Siphonal re- 
tractor muscles straplike, extending ante- 
riorly from the base of the siphons along 
the midportion and over the outer surface 
of the visceral mass to insert on the valves 
just posterior to the umbonal— ventral 
ridge. Ventral adductor muscle, formed as 
a thickened area of the fused mantle mar- 
gin, inserts over and to each side of the 
ventral condyles. Pedal retractor muscles 
pass through the visceral mass and insert 
just anterior to the posterior adductor 
muscle. 

Measurements. See Table 10. I 

Remarks. The young of X. muraokai 
Turner n. sp. superficially resemble those 
of X concava Knudsen, found in the Gulf 
of Panama, but differ in having a mesoplax 
with a large ventral portion, in having the 
excurrent siphon slightly shorter than the 
incurrent siphon, and in having a few large 
cirri on the excurrent siphon. From X. du- 
plicata Knudsen, also from the Gulf of 
Panama, X. muraokai differs in lacking the 
brown periostracal sheath covering the si- 
phons, in having an elongate rather than 
nearly circular posterior adductor scar, and 
in having the dorsal portion of the meso- 
plax reduced, covering only the posterior 
portion of the anterior adductor muscle. 

As with all Xylophaga known to date, 
specimens of X. muraokai show consider- 
able variation in size and scLilpture de- 
pending on the type of wood in which they 
are boring. A specimen boring in green- 
heart with valves only 6 mm in length had 
45 closely spaced denticulated ridges on 



Xylophagainae • Turner 



249 



the anterior slope, whereas a specimen of 
the same size in white pine had only 7 
ridges. Although it is impossible to tell the 
exact age of these specimens, we know 
that the greenheart panel was exposed for 
35 months and the white pine panels were 
exposed for 24 months. Other specimens 
in a white pine panel (exposed on the 
same rack for the same length of time as 
the greenheart panel) reached 12 mm in 
length and had only 24 ridges. 

Large numbers of X. muraokai were 
taken from panels submerged for 2 years 
at 5,640 ft (1,720 m). The openings of the 
burrows averaged about 1 mm in diameter 
and increased rapidly. When specimens 
were uncrowded, the anterior end of bur- 
rows that had reached 15 mm in length 
averaged 12 mm in diameter; burrows 55 
mm long averaged about 15 mm in diam- 
eter anteriorly. The tunnels often ran to- 
gether and frequently two to six specimens 
occupied a large irregular cavity. Appar- 
ently, several specimens occasionally used 
a single opening to the surface. 

Xylophaga muraokai Turner n. sp. is 
known only from the USNCEL Test Site 
I. It has never been taken from panels sub- 
merged at Test Site II, which is just north 
of San Miguel Island in about 2,370 ft (722 
m). Differences in temperature, dissolved 
oxygen content of the water, and hydro- 
static pressure between the two sites are 
probably the responsible factors. It is im- 
possible to say whether or not they are all 
of equal importance. 

Xylophaga washingtona Bartsch, a spe- 
cies of wide geographic and depth range, 
has been taken from wood exposed at Test 
Site I. It is the only species found at Test 
Site II, where it is abundant. For a com- 
parison of the test sites, see Table 11. For 
a description of the test sites, see Muraoka 
(1964, 1965, 1966a, 1966b, 1966c, 1967). 

Range. Known only from USNCEL Test 
Site I. 

Specimens Examined. UNITED STATES, CALI- 
FORNIA: USNCEL Test Site I (33°44'N, 120°45'W) 
about 81 nautical miles SW of Port Hueneme (about 
25 miles S of San Miguel Island, Santa Barbara Is- 



Table II. Comparison of the U.S. Naval Civil 
Engineering Laboratory test sites (average 
values taken from muraoka, 1967). 



Locality 



Deptli 

(fi) 



TeinpeiE 
ture 

(°C) 



Dissolved Hydrostatic 
Ox\'gen Pressure 

(ml/L) (psi) 



Test Site I 
Test Site II 



5,640 
2,370 



2.5 
5.0 



1.26 
.30 



2,482 
1,043 



lands); STU I-l at 5,300 ft (1615.4 m) e.xposed March 
1962-February 1965; STU 1-2 at 5,640 ft (1,720 m) 
exposed October 1963-October 1965; STU 1-4 at 
6,800 ft (2,072.6 m) exposed June 1964-7 July 1965; 
STU 1-5 at 6,000 ft (1,828.8 m) exposed 25 August 
1965-12 Febi-uaiy 1966. 

Xylophaga atlantica Richards 
Plate 20 

Xylophaga atlantica Richards, 1942, Nautilus, 56: 68, 
pi. 6, fig. 4 (east coast of the United States). Ho- 
lotype. Academy of Natural Sciences Philadelphia 
178741; Turner 1955, Johnsonia, 3(34): 152-154, 
pi. 91, figs. 1—6 (type locality. Mount Desert Island, 
Maine). (See Turner, 1955: 153.) [Not Turner, 
1954, Johnsonia, 3(33): 5-6, pi. 4 = X. clenchi 
Turner and Culliney; see below.] 

Distinctive Characters. Mesoplax small, 
anterior to and between the umbos, com- 
posed of two triangular plates that are in 
contact the length of the dorsal inargin 
and meet at a broadly obtuse angle. Ven- 
tral jDortion of the mesoplax narrow, form- 
ing a small posterior cavity into which the 
posterior end of the anterior adductor 
muscle extends. Umbonal— ventral sulcus 
shallow with a median threadlike groove. 
Posterior adductor muscle scar elongate 
and irregularly marked. Excrn-rent siphon 
slightly shorter than the incurrent, the ex- 
current aperture with 15—20 large papillae, 
incurrent aperture with a double row of 
nrmierous minute papillae (Plate 20, Fig. 
3). 

Remarks. Between 16 June 1964 and 16 
July 16 1965, the U.S. NOO submerged 
test panels off Mark Island, Penobscot 
Bay, Maine, in water 200 ft in depth. The 
panels were set in a vertical array at depths 
of 50, 100, 150, and 195 ft. Dissection of 
these panels showed that the bottoin one 
had 85 specimens of X atlantica, the one 



250 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



at 150 ft had six specimens, and the one 
at 100 ft had only one specimen. This sug- 
gests that, hke X. washingtona Bartsch, X 
muraokai Turner n. sp., X. depalmai Turn- 
er n. sp., and probably all Xijlophaga, the 
larvae of this species do not rise very high 
in the water column and that the greatest 
decrease in attack occurs within a few feet 
of the bottom. Probably a panel touching 
or partially submerged in the substrate at 
this locality would have been heavily at- 
tacked. These tests also showed that the 
valves of X. atlantica may reach a length 
of at least 10.5 mm in a year. Several spec- 
imens removed from wood dredged off 
Ipswich, Massachusetts, in about 80 m av- 
eraged 14 mm in length. The maximum 
length for X. atlantica appears to be 
15 mm. 

The Ipswich specimens were well pre- 
served and allow a more detailed descrip- 
tion of the siphons than that given in the 
paper by Turner (1955) cited above. The 
siphons can be extended 1.5—2 times the 
length of the valves. The excurrent siphon 
is slightly shorter than the incurrent si- 
phon and is combined with it for most of 
its length. The excurrent aperture is sur- 
rounded by 15—20 relatively large papillae 
that on the dorsal surface appear to be 
grouped in two lobes. The incurrent si- 
phon is surrounded by an outer rim of 
small papillae and an inner one of slightly 
larger, stouter papillae (Plate 20, Figs. 1, 
2). 

Xylophaga atlantica is oviparous and 
does not brood its young. The reproduc- 
tion and lai'val development of X. atlantica 
are fully discussed by Culliney and Turner 
(1976); they detail inethods of laboratory 
culture and illustrate the various lai"val and 
growth stages. A reexamination of the 
specimen reported by Turner (1954: 5—6, 
pi. 4) has shown it to be X. clenchi Turner 
and Culliney, a species that usually is 
found with young attached to the parent. 
The adult of X. clenchi differs from that of 
X. atlantica in having a mesoplax that lacks 
a ventral portion, in the shape and type of 
marking on the posterior adductor muscle 



scar, in having a broad, rounded ridge pos- 
terior to the umbonal— ventral sulcus, and 
in the arrangement of the papillae on the 
siphons. 

On the basis of the material then avail- 
able, I formerly considered this species 
most closely related to X. washingtona 
Bartsch (Turner, 1956). It is now evident 
that these species belong to two different 
species groups. In X. atlantica, the excur- 
rent siphon is only slightly shorter than the 
incurrent siphon and cirri surround the 
aperture of both siphons (Plate 20, Figs. 
1—3). In addition, the posterior adductor 
scar is irregularly marked, whereas that of 
X. washingtona has regular herringbone 
markings. 

Range. From Newfoundland south to 
Cape Heniy, Virginia, in depths ranging 
from about 15 to 1,242 m. 

Specimens Examined (New Records Since Turner, 
1955). CANADA, NEWFOUNDLAND: Vi miles off 
Argentia in 18.3 m (test panel). NOVA SCOTIA: 
North Bay, Cape Breton Island (46°20'N, 61°50'W) 
in 128 m. UNITED STATES, MAINE: off Mark Is- 
land, Penobscot Bay, in 15.2 m, 30.5 m, 45.7 m, and 
59.5 m (test panels). MASSACHUSETTS: 15 miles 
off Ipswich in 73 m; NE of Cape Cod Light off Truro 
in 40 m; E of Nantucket Island (41°23'N, 68°46'W). 

Xylophaga washingtona Bartsch 
Plate 21 

Xylophaga washingtona Bartsch, 1921, Proceedings 
of the Biological Society of Washington, 34: 32 (San 
Juan Island, Washington). Holotype, U.S. National 
Museum (USNM) 334478; Turner, 1955, Johnson- 
ia, 3(34): 154, pi. 92; Turner, 1956, Nautilus, 70: 
10; Tipper, 1968, Ecological Aspects of Two Wood- 
Boring Molluscs from the Continental Terrace Off 
Oregon, Thesis, Department of Oceanography, 
Oregon State University, pp. 8-13, 64-118. 

Distinctive Characters. Mesoplax small, 
anterior to and between the umbos, com- 
posed of two triangular plates that are in 
contact for the length of the dorsal margin 
and are set at a moderately obtuse angle 
to each other. Ventral portion of the me- 
soplax usually greater than one half the 
width of the dorsal portion and keeled. 
Umbonal-ventral sulcus broad and shal- 
low. Posterior adductor muscle scar elon- 
gate oval in outline with regular herring- 



Xylophagainae • Turner 251 



Table 12. 



Penetration into wood of Xylophaga washingtona at various depths 
and exposure times. 



LocalitN' 



Exposure 



3 miles off Oceanside, California 
STU* II (1 and 2), California 
STU* I (1-1), California 

19 miles off Depoe Bay, Oregon 
25 miles off Depoe Bay, Oregon 
40 miles off Depoe Bay, Oregon 



109 


4 months 


Turner, 1956 




722 


6.5, 13.4 months 


Muraoka, 1965, 


1967 


,615-2,066 


4, 13, 25, and 35 


Muraoka, 1964, 


1966a, 




months 


1966b, 1966c 




200 


38 and 72 days 


Tipper, 1968 




500 


2 months 


Tipper, 1968 




1,000 


2 months 


Tipper, 1968 





STU, submersible test unit. 



bone markings. Proximal end of the coixi- 
bined siphons usually having a thin perios- 
tracal sheath. Excurrent siphon one third 
to one half the length of the incurrent si- 
phon, truncated, and ha\4ng a narrow 
ridge extending from each side for a short 
distance along the dorsal surface of the in- 
current siphon. Excurrent siphonal open- 
ing small, located at the end of a short 
tube extending between the lateral ridges, 
and apparently lacking papillae (Plate 21, 
Figs. 2, 3). Incurrent siphonal opening 
margined by 15 inwardly extending papil- 
lae (Plate 21, Fig. 4). 

Remarks. In recent years a great deal 
has been added to our knowledge of the 
distribution of X. ivashingtona, and inter- 
esting obseivations have been made con- 
cerning its ecology and variation. The 
many specimens taken from the STU pan- 
els exposed by the USNCEL off San Mi- 
guel Island, California, have shown that a 
great deal of variation exists in the shape 
and sculpture of the valves as well as the 
size of the posterior adductor muscle in 
response to the hardness of the substrate 
in which they are boring. This is discussed 
in the section on Variation. From these 
tests we have also learned that lai"val and 
adult X ivashingtona can tolerate a wide 
range in dissolved oxygen concentration. 
They were the only borers taken from the 
STU II test site where the concentration 
was 0.30 ml/L. The temperature at this site 
was 5.0° C (Muraoka, 1965). Based on 
adults dissected from new wood exposed 
at various sites with known dates and 



depth of submergence, it has been possi- 
ble to determine the depth at which the 
larvae can successfully penetrate. These 
data are summarized in Table 12. 

A single large specimen was taken in 
18.3 m from wood that had been removed 
from old street cars and used to make 
Hermosa Reef, V2 mile west of Hennosa 
Beach, Santa Monica Bay, California. This 
is the shallowest record known for the set- 
tlement of lai-vae. The apparent scarcity of 
specimens suggests that this is the upper 
limit of the depth range. Living specimens 
have been taken from a fir log dredged 
from 73 m off Vancouver Island, British 
Colombia. Although no testing has been 
done in this area, it is likely that, in these 
colder waters, X. washingtona could occur 
at shallower depths. 

Muraoka (1966c) found that "wood 
specimens which were exposed near the 
sediment were damaged considerably 
more severely than those specimens which 
were exposed about 3 feet above the sed- 
iment." This, he said, "indicates that the 
deep sea borers are very active in large 
numbers immediately above the sediment 
layer and that their numbers tend to de- 
crease in seawater as the distance from the 
sediment layer increases." Tipper (1968) 
showed that the settlement of lai-vae of X. 
ivashingtona off Depoe Bay, Oregon, was 
densest in proximity to the sea— sediment 
interface and that the drop in borer pen- 
etration usually occurred within the first 6 
cm upward from the interface. He also 
showed that, from the initial penetration 



252 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



very close to the sea— sediment interface, 
the attack progressed upward with increas- 
ing time of exposure. Analyses of these 
data suggest that 1) the free-swimming lar- 
vae of X. washingtona probably do not rise 
more than 10-25 ft above the sea floor; 2) 
as the attack increases, competition occurs 
for space; and 3) the second crop of larvae, 
prevented from settling on the lower levels 
because of the activity of the siphons of 
specimens already in the wood, are kept 
swimming and eventually settle on surfac- 
es higher up in the water column. 

Tipper (1968) also pointed out that the 
depth of penetration decreased with in- 
creased density of the wood. Cedar, pine, 
fir, and oak panels exposed for 50 days at 
a depth of 200 m showed an average pen- 
etration of 3.8 mm in cedar, 3 mm in pine, 
1.5 mm in fir, and 0.5 mm in oak. De- 
crease in depth of penetration also is re- 
lated to increase in depth of exposure. 
This, he postulated, may be correlated 
with lower temperatures resulting in slow- 
er growth rates or with the increased den- 
sity of the wood from compression at great 
depth. 

These obseivations agree with those ob- 
tained from the tests off southern Califor- 
nia (see section on Variation in the Intro- 
duction). From the STU samples we found 
that the softer the wood (except balsa), the 
deeper the Xylophaga penetrated and the 
greater was the development of the chim- 
ney of fecal pellets. In a cedar panel ex- 
posed on STU 1-5 for 6 months, two sets 
of X washingtona were found; the earlier 
set had tunnels averaging 10—12 mm in 
length, and the second set had tunnels av- 
eraging 5 mm in length. Another panel on 
STU I-l exposed for 35 months had spec- 
imens with tunnels reaching 35 mm in 
length. These tunnels averaged 10 mm in 
diameter at the anterior end; a chimney of 
rather coarse fecal pellets lined the pos- 
terior 15 mm of the burrow. Considering 
the greater depth of the STU panels and 
realizing that the rate of growth decreases 
with age, these growth rates agree with 
those reported by Tipper (1968). 



Based on data gathered to date from 
test panels, it would appear that X. wash- 
ingtona probably breeds throughout the 
year and that the entire larval life is spent 
in the sea. Young specimens have never 
been found attached to the shells of the 
hundreds of adults examined. 

Range. From Vancouver Island, British 
Columbia, south to the Santa Barbara Is- 
lands, California, in depths ranging from 
20 to 2,000 m. 

Specimens Examined (New Records Since Turner, 
1955). CANADA, BRITISH COLOMBIA: Satellite 
Channel, between Saltspring and Vancouver Islands 
in 93 m (C. Carl). UNITED STATES, OREGON: 
about 30 miles W of Seaside (46°00'N, 124°48'W) in I 
464.5 m; about 50 miles W of Tillamookhead 
(45°54'N, 125°09'W) in 1,554.4 m; about 40 miles W 
of Silver Point (45°52'N, 124°54'W) in 822.9 m (all 
Oregon State University); 19 miles W of Depoe Bay 
(44°52'N, 124°54'W) in 200 m; 25 miles W of Depoe 
Bay (44°52'N, 124°36'W) in 500 m; 40 miles W of 
Depoe Bay (44°52'N, 125°01'W) in 1,000 m (all R. 
Tipper). CALIFORNIA: Hermosa Reef, Vi mile off ' 
Hermosa Beach in 18.3 m (J. Fitch); 3 miles off : 
Camp Pendleton pier, Oceanside, in 100 m (F. Snod- . 
grass); USNCEL STU II (1 and 2), 75 miles W of: 
Port Hueneme or about 5 mi NW of San Miguel I., 
Santa Barbara Islands (34°06'N, 120°42'W) in 722 ni; 
USNCEL STU I (1, 4, and 5), 81 miles SW of Port 
Hueneme or 25 miles SW of San Miguel Island, San- 
ta Barbara Islands (33°44'N, 120°45'W) in 1,524- 
2,066 m (both J. Muraoka); Allan Hancock station 
1372—41, about % mile E of Empire Landing, Santa 
Catalina Island, Santa Barbara Islands (.3.3°25'50"N, 
118°24'50'^V) in 84 m (Allan Hancock Foundation); 
2 miles off Eel Point, San Clemente Island, Santa 
Barbara Islands in 72.2 m (F. Snodgrass). 

Xylophaga rikuzenica Taki and Habe 
Plate 22 

Xylophaga rikuzenica Taki and Habe, 1945, Japanese 
Journal of Malacology (Venus), 14: 112 (off Riku- 
zen, Honshu, Japan). Holotype, T Habe Collec- 
tion; paratype, MCZ 194821. Neoxylophaga riku- 
zenica (Taki and Habe) 1950, Illustrated Catalogue 
of Japanese Shells, no. 7, p. 46, text-figs. 4, 5. 

Distinctive Characters. Mesoplax com- 
posed of two inflated triangular plates with 
a large basal portion and ventral keel; 
plates in contact both dorsally and ven- 
trally for the length of the medial margin 
(Plate 22, Figs. 5-7). Posterior adductor 
muscle scar elongate oval, tapering dorsal- 



Xylophagainae • Turner 



253 



ly, and with regular herringbone markings. 
Valves and dorsal plates covered with a 
heavy, brown periostracum. Umbonal- 
ventral sulcus wide and deep (Plate 22, 
Figs. 2, 4). 

Remarks. Taki and Habe (1945) briefly 
described this species in Japanese but in- 
cluded no figures. In 1950, they created 
the genus Neoxylophaga with rikuzenica as 
the type species, mentioning the small, tri- 
angular mesoplax and thick, brown perios- 
tracum as generic characters. Turner 
(1956) related X rikuzenica to X. wash- 
ingtona Bartsch on the basis of the dorsal 
plates. A paratype specimen, received 
through the kindness of Dr. Habe, shows 
that the posterior adductor muscle scar is 
similar to that of X washingtona Bartsch, 
X praestans Smith, and X. aiirita Knud- 
sen, the other species in this group with 
simple, unfringed lappets extending from 
the truncation of the excurrent siphon. Xy- 
lophaga rikuzenica is closely related to but 
differs from X washingtona mainly in hav- 
ing a heavy, brown periostracum covering 
the valves and mesoplax as well as in hav- 
ing a much wider, deeper umbonal— ventral 
sulcus. Unfortunately, the siphons of X ri- 
kuzenica are unknown so comparisons 
cannot be made on that basis. However, 
all species with regular herringbone mark- 
ings on the posterior adductor scar known 
to date have similar siphons so X. rikuzen- 
ica can be assumed to belong to the same 
group. No Xylophaga have been taken 
north of Vancouver, British Columbia, but 
continuous collecting across the north Pa- 
cific may show that X. rikuzenica merges 
with X. washingtona. Until this is done, 
the two species are being maintained be- 
cause they are recognizable and may not 
merge. 

Range. Known only from off Rikuzen 
and Toyama Bay, Honshu, Japan, in depths 
ranging from 100 to 700 fathoms (183 to 
1,270 m) (Taki and Habe, 1950). 



Specimens Examined. JAPAN: off Rikuzen, Hon- 
shu in 183 m. 



Xylophaga depalmai*^ Turner new species 
Plates 23, 24 

Holotype. MCZ 316735. 

Type Locality. 3.2 miles off Fort Lau- 
derdale, Florida (26°04'N, 80°04'W) in 
152.4 m (500 ft) in a test panel. 

Distinctive Characters. Shell globose, 
anterior slope with fine, closely set, den- 
ticulated ridges; umbonal— ventral sulcus 
shallow and bounded posteriorly by an in- 
distinct ridge; posterior slope smooth. 
Umbonal— ventral ridge narrow and high. 
Muscle scars barely impressed and 
smooth. Mesoplax composed of two inflat- 
ed, elongate plates that are coiled poste- 
riorly, vary in shape, and are occasionally 
fused. Excurrent siphon truncate; lappets 
on dorsal surface of incurrent siphon with 
exceedingly fine serrations. 

Description. Shell globose, valves reach- 
ing 9.8 mm in length and 9 mm in height, 
thin, fragile, and with a thin transparent 
periostracum. Pedal gape angle about 95- 
105°. Anterior slope sculptured with ex- 
ceedingly fine, closely set, denticulated 
ridges; specimens 8 mm long had 40—66 
ridges. Posterior portion of the anterior 
slope rather wide and the ridges extending 
to the umbonal— ventral sulcus more 
coarsely denticulate. Umbonal— ventral sul- 
cus barely impressed, nearly smooth, and 
bounded posteriorly by a fine, indistinct 
ridge. Disc and posterior slope faintly 
marked with growth lines. Posterior slope 
with a low, smoothly cui"ved dorsal margin 
that is not reflected. 

Inner surface of valves smooth and glis- 
tening. Umbonal-ventral ridge narrow, 
high, and not enlarging at the ventral mar- 
gin to form a knoblike condyle. Posterior 
adductor muscle scar barely impressed, 
rather small, oval, smooth, and set high on 
the posterior slope. Pedal retractor scar 
dumbbell-shaped or as two adjacent cir- 



'^ Named for Jolin DePalma, U.S. Naval Oceano- 
grapliic Office, wlio was responsible for the tests off 
Fort Lauderdale, Florida; in the Tongue of the 
Ocean, Bahama Islands; and in Penobscot Bay, 
Maine. 



254 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 





Table 13. Measurements 


OF 






Xylophaga depalmai. 




Length 


Height 






(mm) 


(mm) 


Locati(jn 




6.0 


5.5 


off Fort Lauderdale, 


Florida 


7.6 


7.0 


off Fort Lauderdale, 


Florida 


8.0 


7.5 


holotype 




8.2 


8.2 


off Fort Lauderdale, 


Florida 


8.5 


7.0 


off Fort Lauderdale, 


Florida 


9.8 


9.0 


off Fort Lauderdale, 


Florida 



cles just anterior to the posterior adductor 
scar. Ventral adductor scar large, elongate, 
irregLilarly oval, located posterior to the 
umbonal— ventral ridge and paralleling the 
ventral margin of the valve. Siphonal re- 
tractor scar small, broadly oval, and locat- 
ed just posterior to the umbonal-ventral 
ridge at the level of the pedal retractor 
scar. Anterior adductor scar covering the 
umbonal reflection and the lower surface 
of the mesoplax. Chondrophore and inter- 
nal ligament well developed. 

Mesoplax composed of two inflated, 
posteriorly coiled plates that may be asym- 
metrical, have a well-developed ventral 
portion, are longer than wide, with the 
medial margin of the plates parallel and 
occasionally with the two parts fused 
(Plate 24, Figs. 3-17). 

Burrow length of specimens with valves 
4—8 mm in length varying from 1.5 to 4 
times the length of the valves. Posterior 
end of the burrows usually with a smooth, 
firmly packed chimney composed of fine 
wood fragments. 

Siphons capable of complete retraction 
within the valves. Excurrent siphon trun- 
cated just posterior to the valves and lack- 
ing cirri. Finely serrated lappets extend 
from the truncation along either side of 
the dorsal sru'face of the incurrent siphon. 
Incurrent siphon with fine cirri surround- 
ing the aperture. Base of the siphons cov- 
ered by a fine periostracal sheath that ex- 
tends along the sides of the siphons and 
contains fine, irregular, glasslike granules 
(Plate 23, Figs. 1, 4, 5). 

Measurements. See Table 13. 



Remarks. This species is most closely re- 
lated to X. guineensis Knudsen from the 
Gulf of Guinea, West Africa. Xylophaga 
depalmai differs in being larger, having an 
elongate mesoplax, the posterior end of 
which fits between the umbos, and in hav- 
ing more coarsely serrated lappets on the 
siphons. (See also Remarks under X gui- 
neensis.) From X. tipperi Turner n. sp., to 
which it is also related, X. depalmai differs 
in having an elongate inflated mesoplax 
that fits between the umbos rather than a 
broad flattened one that covers them, in 
having finely serrated lappets on the si- 
phons, in having the granules on the si- 
phons only in the periostracum rather than 
being embedded in the tissue, and in pro- 
ducing a chimney. From X. mexicana Dall,l 
which it also somewhat resembles, X. de- 
palmai differs in the shape of the meso- 
plax, in having narrower, more finely 
fringed lappets on the siphons, an incon- 
spicuous ridge posterior to the umbonal- 
ventral sulcus, and an exceedingly thin 
periostracum. In addition, the granular in- 
clusions on the siphons of X mexicana are 
coarse, chalky, white, and arranged in a, 
single line extending nearly the length of 
the siphons. 

Variation exhibited in the mesoplax of X. 
depalmai is unusual and unexpected, for 
in most species of Xylophaga the mesoplax 
is remarkably uniform vmless the specimen 
is injured or stenomoiphic. The extreme 
forms (Plate 24, Figs. 11-15) are suffi- 
ciently different to be considered of spe- 
cific value in this genus. However, in the 
more than 300 specimens obtained from 
the test site off Fort Lauderdale, connect- 
ing forms exist between them. The char- 
acters of the shell, the siphon, and soft' 
parts are similar in all specimens. Conse- 
quently, because the large majority are 
typical (Plate 24, Figs. 16, 17), the ex-. 
tremes are considered to be variants. 

Xylophaga depalmai Tm^ner n. sp. was 
the most common species occm-ing in the 
Fort Lauderdale test. It ranged from 1 to 
3.2 miles off shore and occurred in boards 
placed near the bottom in depths from 100 



Xyloph AGAIN AE • Turner 



255 



Table 14. Data for all of the specimens of Xylophaga depalmai n. sp. found in the collecting 

panels off fort lauderdale, florida. 





Distance 


















From 


Bottom 


Dentil ol 










No. 




Shore 


Deptli 


Panel 


Date 


ol 


Dati 


of 


.Sj5eeimens 
Examined 


Panel no. 


(miles) 


in Feet (i 


i) in Feet (m) 


Subniei 


genee 


Rem 


jval 


2 panels without 


2.5 


300 


on bottom 


Sept. 1961 


June 1962 


±250 


numbers 




(91.44 














W44 


1.8 


100 


on bottom 


Jan. 1964 


Oct. 1964 


1 






(30.48 














H50 


2.3 


200 


100 


Jan. 1964 


Oct. 1964 


2 






(60.96 


(30.48) 












H56 


2.3 


200 


on bottom 


Jan. 1964 


Sept. 1964 


5 






(60.96 














G62 


2.6 


300 


200 


Jan. 1964 


Sept. 1964 


6 






(91.44 


(60.96) 












G68 


2.6 


300 


on bottom 


Jan. 1964 


Sept. 1964 


31 






(91.44 














J59 


2.3 


200 


on bottom 


24 Jan. 


1964 


4 Jan. 1965 


3 






(60.96 














492 


3.2 


500 
(152.4 


200 
(60.96) 


20 Oct. 


1965 


13 Oct. 


1965 


3 


493 


3.2 


500 
(152.4 


200 
(60.96) 


20 Oct. 


1965 


13 Oct. 


1966 


10 


497 


3.2 


500 

(152.4 


300 

(91.44) 


20 Oct. 


1965 


13 Oct. 


1966 


5 


498 


3.2 


500 

(152.4 


300 

(91.44) 


20 Oct. 


1965 


13 Oct. 


1966 


7 


500 


3.2 


500 
(152.4 


400 

(121.92) 


20 Oct. 


1965 


13 Oct. 


1966 


3 


501 


3.2 


500 
(152.4 


400 
(121.92) 


20 Oct. 


1965 


13 Oct. 


1966 


2 


502 


3.2 


500 

(152.4 


400 

(121.92) 


20 Oct. 


1965 


13 Oct. 


1966 


3 


504 


3.2 


500 

(152.4 


on bottom 


20 Oct. 


1965 


13 Oct. 


1966 


8 


505 


3.2 


500 

(152.4 


495 


20 Oct. 


1965 


13 Oct. 


1966 


6 



to 500 ft (30.5 to 152.4 m). In addition, a 
few specimens were found in boards 100, 
200, and even 300 ft off the bottom, which 
means that, in this species at least, the lar- 
vae range well up in the water column. As 
can be seen from Table 14, panels on the 
bottom in 300 ft (91.4 m) were the most 
heavily attacked. This may reflect cyclic 
variation in population density because the 
panels were exposed at different times, or 
that, because of the configuration of the 
bottom and the currents, inore wood was 
available on the bottom in this vicinity, 
supporting a native population and a ready 
source of larvae. However, this also may 



indicate that this is the optimum depth for 
this species. 

According to DePalma (1969), the salin- 
ity at the test site was uniformly high and 
the temperature of the water ranged from 
24 to 30° C at the surface, from 20 to 25° 
C at 100 m, and from 7 to 9.5° C at 153 
in. The northward-flowing Florida Current 
averaged 1.5 knots. 

Range. From Florida north to Massa- 
chusetts in depths ranging from about 30 
to 174 m. 

Specimens Examined. UNITED STATES, FLOR- 
IDA: off Fort Lauderdale (26°04'N, 80°04'W). See 
Table 14. Additional records include: FLORIDA: 160 



256 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



miles W of Tampa, OTEC station 3089 in 126 ni; and 
OTEC station 3086 in 96 m. RHODE ISLAND: 
From rosewood panels tied to lobster pot set at 
39°57'N, 69°19'W in 45.7 m (Knutton, from Al Ea- 
gles ship Reliance). MASSACHUSETTS: Delaware 
II, station 369 (42°37'N, 66°27'W) from wood 
dredged from 174 m. 

Xylophaga guineensis Knudsen 
Plates 25, 26 

Xylopliaga guineensis Knudsen, 1961, Galathea Re- 
port, 5: 195-196, fig. 38 (Galathea, Station 52, off 
West Africa, 01°42'N, 07°51'E in 2,550 m). Holo- 
type. Zoological Museum, University of Copenha- 
gen. 

Distinctive Characters. Valves globose, 
with exceedingly fine, closely spaced, den- 
ticulated ridges; umbonal— ventral sulcus 
barely impressed, bounded by a weak an- 
terior ridge and a well-marked, narrow 
posterior ridge. Mesoplax composed of 
two inflated cornucopialike plates. 

Description. Valves globose, reaching 
2.2 mm in length, thin, and fragile. Pedal 
gape angle about 90°. Anterior slope sculp- 
tured with exceedingly fine, evenly spaced, 
denticulated ridges, there being about 45 
in a specimen 2 inm long. Umbonal— ven- 
tral sulcus only slightly impressed, bound- 
ed anteriorly by a low, inconspicuous, 
rounded ridge and posteriorly by a narrow, 
well-defined ridge, occasionally almost 
bladelike. Disc and posterior slope 
smooth. 

Inner surface of valves smooth and glis- 
tening. Muscle scars barely visible. Poste- 
rior adductor scar smooth, lobed anteriorly 
with the small pedal retractor just anterior 
to it in the upper embayment. Umbonal— 
ventral ridge and condyle prominent, with 
a parallel groove posterior to the ridge. 
This groove is the internal expression of 
the ridge bounding the sulcus on the outer 
surface. 

Mesoplax composed of two smooth, in- 
flated, cornucopialike plates, the horns of 
which curl toward each other (Plate 25, 
Figs. 3, 6, 8, 9); the two halves usually 
equal and mirror images but occasionally 
unequal and varying in size and shape. 

Siphons about equal to the length of the 





Table 15. Measurements of 






Xylophaga guineensis. 


Length 


Height 




(mm) 


(mm) 


Location 


0.9 




Pillsbury, station 260 


1.3 


1.2 


Pillsbunj, station 260 


1.3 


1.5 


paratype (estimated from Knudsen, 
1961, fig. 38b) 


1.5 


1.3 


Atlantiqiie Sud, station 33 


2.2 


2.2 


Atlantique Sud, station 33 



shell, capable of complete retraction with- 
in the valves. Excurrent siphon truncated, 
about one third the length of the incurrent 
siphon, with narrow and minutely serrated 
lappets extending from the truncation 
along the dorsal surface of the incurrent , 
siphon. Base of siphons covered by a filmy' 
periostracal sheath with fine, irregular, 
whitish granules imbedded in it along the 
sides of the siphons (Plate 25, Figs. 1, 2). 

Measurements. See Table 15. 

Remarks. When Knudsen described X 
guineensis he had only "very fragmented 
shells of 5—6 individuals removed froin a 
piece of wood; not a single complete shell I 
present." One specimen had a complete 
mesoplax although only the umbonal area '■ 
of the valves was present. The species was 
diagnosed on the basis of the ixiesoplaxi 
and the large sculptured, bright yellow i 
prodissoconch. 

In the course of examining wood 
dredged from off the western coast of Af- 
rica, I obtained four dead specimens, one 
with a complete mesoplax {Atlantique Sud, 
station 33), and seven small living speci- 
inens, two complete {Pillsbury, station 
260), all of which appear to be this species. 
The two plates of the mesoplax of these 
specimens are mirror images of each other 
(Plate 26, Figs. 2-4). As shown under X. 
depalmai (Plate 24, Figs. 11, 12), the me- 
soplax is not always bilaterally syinmetrical. 
Therefore, these new specimens from off 
the western coast of Africa are being con- 
sidered X. guineensis and a description of 
the species based on entire specimens is 
given. 



Xyloph AGAIN AE • Turner 



257 



Although stenomorphs are commonly 
found in Xijlophaga (see section on Vari- 
ation), the specimens dissected from the 
wood dredged by the Atlantique Sud Ex- 
pedition and the Fillsbunj did not appear 
to be stunted or malformed in any way 
They were not crowded and the wood was 
not particularly hard. Consequently, on the 
basis of the large number of evenly spaced 
ridges on the anterior slope, the well-de- 
veloped inesoplax, the strength of the 
ridge bounding the umbonal— ventral sul- 
cus posteriorly, and the small size of the 
specimens, I consider this to be a small 
species. 

All the major characters of the siphons 
mentioned in the description could be 
seen with a dissection scope at 250 X. 
However, the fringe on the lappets could 
only be detected under a compound mi- 
croscope at 430 X. It was impossible to de- 
termine whether or not the fringe extend- 
ed the entire length of the lappet on the 
two specimens that had extended siphons. 
The siphons appear similar to those of X 
depalmai. 

Xylophaga guineensis is most closely re- 
lated to X. depalmai taken from waters off 
Florida and the Bahamas. Xijlophaga gui- 
neensis differs in being smaller; in having 
a smooth mesoplax composed of inflated, 
tubular, cornucopialike plates; in having 
more numerous and closely spaced, den- 
ticulated ridges on the anterior slope when 
comparing specimens of equal size; and in 
having a larger, more prominent, strongly 
sculptured prodissoconch. (See also Re- 
marks under X. depalmai. ) 

The specimens that Knudsen described 
were taken from wood dredged in 2,550 m 
but the three lots of new material were 
from 147, 145, and 46 m. Although X. gui- 
neensis may extend into deep water, it is 
probably a fairly shallow water species, 
and this would agree with the known 
depth range of X depalmai, the species to 
which it is most closely related. The fact 
that Knudsen's specimens were all dead 
and fragmented further substantiates the 
possibility that the wood was carried into 



deeper water after it was invaded by X. 
guineensis. The continental shelf in this 
area is narrow and such a movement of 
wood on the bottom could easily take 
place. 

Range. Gulf of Guinea, West Africa, in 
depths ranging from 46 to 147 m (living 
material) and in 2,550 m (dead). 

Specimens Examined. CAMEROONS: Pilhhitrtj, 
station 260, off Santa Isabel Island (3°45'N, 9°05'E) 
in 46 m. GABON: Atlantique Sud, station 146, about 
46 miles NNE of Port Gentil (0°0.3'S, 9°07'E) in 147 
m; Atlantique Sud, station 33, 35 miles W of Ambri- 
zette (7°16'S, 12°17'E) in 145 m. 

Xylophaga mexicana Dal I 
Plates 27, 28 

Xylophaga mexicana Dall, 1908, Bulletin Museum of 
Comparative Zoology, 43(6): 425 {Albatross, station 
3422, off Acapulco, Mexico [16°47'N, 99°59'W] in 
141 fathoms [257.9 m]). Holotype USNM 122947; 
Turner, 1955, Johnsonia, 3(34): 150, pi. 90. 

Distinctive Characters. Umbonal— ven- 
tral sulcus bounded posteriorly by a nar- 
row, shaij) ridge. Mesoplax ear-shaped, in- 
flated, the two halves not meeting medially 
except at the posterior end. Excurrent si- 
phon truncated; lappets extending along 
the dorsal surface of the incurrent siphon 
finely fringed. Siphons with a single row of 
opaque white granules imbedded along 
the sides. Posterior quarter of the incur- 
rent siphon papillose. 

Description. Shell globose, valves reach- 
ing 10.5 mm in length and 9.5 mm in 
height, thin, fragile, and with a thin, light 
straw-colored periostracum on the poste- 
rior slope. Pedal gape angle about 110°. 
Anterior slope with exceedingly fine, close- 
ly set, denticulated ridges. Umbonal-ven- 
tral sulcus rather narrow, moderately deep, 
and bounded posteriorly by a narrow, 
sharp ridge. Disc and posterior slope 
sculptured with fine growth lines. Poste- 
rior margin broadly rounded. Posterior 
slope becoming proportionately more 
elongate with increased size and age, as 
indicated in the measurements. Umbonal 
reflection free and recui-ved anteriorly, ap- 



258 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



pressed posteriorly, and covering the an- 
terior portion of the umbonal area. 

Inner surface of the valves white, 
smooth, and glazed. Umbonal-ventral 
ridge narrow, slightly segmented, and 
broadening only slightly toward the ventral 
margin to form the condyle. Posterior ad- 
ductor muscle scar well impressed, 
smooth, broadly oval in outline, set high 
on the posterior slope and at the posterior 
margin of the valve. Pedal retractor scars 
nearly circular, double, and adjacent to the 
embayment on the anterior margin of the 
posterior adductor scar. Ventral adductor 
scar large, irregularly oval in outline, and 
well posterior to the umbonal-ventral 
ridge. Siphonal retractor scar small, bi- 
lobed, and located posterior to the um- 
bonal-ventral ridge at a level correspond- 
ing to the pedal retractor scars. Anterior 
adductor scar covering the umbonal re- 
flection. Chondrophore and internal liga- 
ment well developed. 

Mesoplax somewhat variable but usually 
slightly longer than wide, with the medial 
margin of the two halves diverging. Plates 
ear-shaped, inflated, coiled posteriorly, and 
covering the umbos. 

Siphons capable of complete retraction 
within the valves and of extension to at 
least 1.5 times the length of the valves. Ex- 
current siphon truncated, about one third 
the length of the incurrent siphon, and 
lacking cirri. Lappets extending from the 
truncation of the excurrent siphon finely 
fringed. Posterior portion of the incurrent 
siphon papillose; inner edge of siphonal 
opening margined with numerous minute 
cirri. Siphons with thin periostracal sheath 
posteriorly and with a single row of white, 
opaque, irregular granules embedded 
along the sides. 

Measurements. See Table 16. 

Remarks. Xylophaga mexicana Dall was 
described on the basis of two right valves 
that were in rather poor condition and 
lacked a mesoplax. Consequently, it has 
been impossible up to now to relate it to 
other species. Valves of living specimens 
taken off California agree with those of the 



Table 16. Measurements of 

Xylophaga mexicana. 



Length 


Height 




(mm) 


(mm) 


Locatioi 1 


4.8 


4.5 


Malibu Reef, California 


5.0 


4.2 


holotype 


7.5 


7.3 


Malibu Reef, California 


8.0 


7.3 


Santa Monica Reef, California 


9.0 


9.5 


Hermosa Reef, California 


10.5 


9.0 


Santa Monica Reef, California 



holotype of X. mexicana. Dall's specimen 
was from much deeper water (141 fath- 
oms) than the Californian series (about 19 
fathoms) but it could have been advecti- 
tious at that depth. Xylophaga mexicana 
Dall could, of course, be considered a no- 
men duhium but this would necessitate de- 
scribing the Californian specimens. Be- 
cause the valves of these specimens resem- 
ble the holotype of X mexicana sufficiently 
well to carry the name, it seems best to 
establish that taxon firmly on the basis of 
this new material. 

The size of the beaks, the smooth mus- 
cle scars, and the narrow, rather deep um- 
bonal-ventral sulcus with a pronounced 
ridge at its posterior margin relate X mex- 
icana to Xylophaga glohosa Sowerby from 
Chile (Turner, 1955). Xylophaga mexicana 
differs from X. globosa in having a much 
finer fringe on the lappets, in having a se- 
ries of irregular, opaque granules along the 
sides of the siphons, and in having the pos- 
terior portion of the incurrent siphon pa- 
pillose. In addition, the mesoplax of X. 
mexicana is proportionately much smaller 
than that of X. globosa, and the two plates 
diverge medially. In his original descrip- 
tion, Dall (1908) did not relate X. mexi- 
cana to any other species in the genus. 
One year later (Dafl, 1909), he gave the 
range of X. globosa Sowerby as from Pan- 
ama to Chile but cited no specific locali- 
ties. The Panamanian specimens on which 
he based this range may well have been X. 
mexicana. I have not seen a specimen of 
X. glohosa from north of Chiloe Island, 
Chile. The valves of X. cluplicata some- 



Xylophagainae • Turner 



259 



what resemble those of X rnexicana; how- 
ever, the siphons of these two species 
place them in different groups. (See also 
Remarks under X. tipperi Turner n. sp. 
and X japonica, the species to which X. 
rnexicana is most closely related.) 

Through the kindness of John Fitch and 
Charles Turner of the California State 
Fisheries Laboratory, Terminal Island, 
California, I received 55 specimens of X. 
rnexicana taken from test panels exposed 
on the "replication reef experiments" con- 
ducted by that laboratory. Old streetcars, 
automobile bodies, concrete shelters, and 
rocks were dumped offshore in several lo- 
calities to sei've as settling areas for marine 
organisms and to make the area more at- 
tractive to fish. Within a short time, the 
test panels and wooden portions of the 
streetcars were attacked by teredinids and 
Xijlophaga. Because all the specimens had 
been dissected from the wood, it is im- 
possible to say whether or not they pro- 
duced a chimney 

Range. Living specimens known only 
from Santa Monica Bay, California, in 
about 35 m. 

Specimens Examined. MEXICO: Bocochibampo, 
Sinaloa (dead). UNITED STATES, CALIFORNIA: 
All from "replication reefs" off Hermosa, Malibu, Re- 
dondo, and Santa Monica (about 33°50'N, 118°30'W) 
in about 35 m. 

Xylophaga tipperi Turnev new species 
Plate 29 

Holotijpe. MCZ 316736; paratype, MCZ 
316737. 

Type Locality. 3.2 miles off Fort Lau- 
derdale, Florida (26°04'N, 80°04'W) in 
152.4 m (500 ft) in a U.S. NOO test panel, 
submerged from October 1965 to October 
1966. 

Distinctive Characters. Umbonal— ven- 
tral sulcus bounded by low rounded ridg- 
es. Mesoplax ear-shaped, slightly longer 
than wide, compressed, and with a sharp 
peripheral keel. Excurrent siphon truncat- 
ed; lappets on dorsal surface of incurrent 
siphon with a coarse fringe. Siphons with 
a single row of minute glasslike plaques 



embedded along the side and with knobby 
pustules at the posterior end. 

Description. Shell globose, reaching 9 
miTi in length and 8.5 mm in height, thin, 
fragile, with prominent umbos and a dull, 
light brown periostracum. Pedal gape an- 
gle about 110°. Anterior slope sculptured 
with numerous close-set, denticulated 
ridges, there being 25 on the holotype. 
Umbonal— ventral sulcus slightly de- 
pressed, bounded by a threadlike rounded 
ridge anteriorly and a somewhat heavier 
one posteriorly. Posterior slope low and 
sculptured with distinct growth lines. Um- 
bonal reflection free for most of its length, 
the ventral edge of the mesoplax fitting be- 
neath it anteriorly. 

Inner surface of the valves smooth and 
glistening. Umbonal— ventral ridge very 
prominent, nearly sinooth, and not greatly 
enlarged ventrally at the condyle. Chon- 
drophore and internal ligament well de- 
veloped. Posterior adductor muscle scar 
oval, tapering dorsally, only slightly im- 
pressed, and smooth. Pedal retractor scar 
kidney-shaped and located adjacent to the 
anterior margin of the posterior adductor 
scar, about midway dorsoventrally. Siphon- 
al retractor scar lightly impressed, located 
just posterior to the umbonal— ventral ridge 
at the level of the ventral margin of the 
beak. Ventral adductor scar usually not vis- 
ible but located near the ventral margin 
posterior to the umbonal— ventral ridge. 

Mesoplax large, ear-shaped, covering 
the umbos, longer than wide, compressed 
dorsoventrally, with a shaq? peripheral 
keel, and a coiled early portion. 

Siphons united, excurrent siphon trun- 
cated, about one third the length of the 
incurrent siphon. Lappets extending from 
the truncation along the dorsal surface of 
the incurrent siphon with a coarse fringe. 
Incurrent siphon papillose posteriorly, 
with a single row of glasslike plaques im- 
bedded along the side at the juncture of 
the lappets and the excurrent siphon and 
with numerous small, broad papillae sur- 
rounding the inner rim of the aperture. 



260 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 17. Measurements of Xylophaga tipperi. 



Lens;tli 


Height 






(iTini) 


(mm) 


Location 




2.0 


2.0 


off Fort Lauderdale, 


Florida 


4.6 


4.1 


off Fort Lauderdale, 


Florida 


7.5 


7.0 


off Fort Lauderdale, 


Florida 


8.2 


7.5 


off Fort Lauderdale, 


Florida 


9.0 




holotype 





Burrows reaching 20 mm in length. Fe- 
cal pellets not formed into a chimney. 

Measurements. See Table 17. 

Remarks. This species is closely related 
to X. mexicana Dall from California but 
differs in having a compressed, sharply 
keeled mesoplax; in having glasslike rather 
than white, chalky plaques imbedded in 
the siphons; and in having much more 
coarsely fringed lappets on the siphons. 
Xylophaga tipperi differs from Xylophaga 
dorsalis Turton in having a definitely 
sculptured mesoplax, an excurrent siphon 
one third to one half the length of the in- 
current (in X. dorsalis it is truncated at the 
posterior end of the valves), and in having 
glasslike plaques imbedded in the walls of 
the siphons. In addition, on the basis of 
the limited material now available, X. tip- 
peri apparently does not build a chimney. 
Xylophaga tipperi differs from X. depahnai 
Turner n. sp., also from off Fort Lauder- 
dale, in having a compressed, sharply 
keeled mesoplax, a much coarser fringe on 
the lappets, and in having glasslike plaques 
imbedded in a single line along the sides 
of the siphons. In addition, the posterior 
end of the incru-rent siphon of X. tipperi 
is papillose. Xylophaga hayeri Turner n. 
sp., which also is found in Florida and the 
West Indies, differs from X. tipperi and all 
other species in Group 6 in having a 
broad, horizontal mesoplax (Plate 31, Fig. 
3). 

Only eight specimens of X tipperi Turn- 
er n. sp. were obtained. They were all 
from panel 505 and were collected along 
with X. depahnai Turner n. sp. The spec- 
imens, even the smallest, were consistent 
in having a broad, compressed mesoplax as 



well as in the characteristics of the si- 
phons. For data on the testing site, see in- 
formation under X. depahnai. 

Range. Known only from off Fort Lau- 
derdale, Florida, in 152.4 m. 

Specimens Examined. UNITED STATES, FLOR- 
IDA: 3.2 miles off Fort Lauderdale (26°04'N, 
80°04'W) in 152.4 m (500 ft) in a test panel. 

Xylophaga bayerr Turner new species 
Plates 30, 31 

Holotype. MCZ 316738; paratype, MCZ 
316739. 

Type Locality. U.S. NOO test site, about 
3.2 miles off Fort Lauderdale, Florida 
(26°04'N, 80°04'W) in 152.4 m (500 ft). 
From test panels submerged froixi October 
1965 to October 1966. 

Distinctive Characters. Posterior margin 
of the umbonal-ventral sulcus bounded by 
a pronounced, sharp ridge. Mesoplax of 
adult much broader than long, extending 
laterally as wings and with a sharp periph- 
eral margin. Excurrent siphon truncated, 
about one third the length of the incurrent 
siphon. Lappets extending along the dorsal 
surface of incurrent siphon finely fringed. 

Description. Shell globose, reaching 8 
mm in length, thin, fragile, with prominent 
umbos and a heavy, golden-brown perios- 
tracum covering the shell and the meso- 
plax. Pedal gape angle about 120°. Beaked 
portion of anterior slope sculptured with 
numerous, closely set, denticulated ridges, 
there being 32 on the holotype. Posterior 
portion of the anterior slope narrow, the 
ridges very closely packed. Umbonal re- 
flection broad, closely appressed over the 
umbos, free anteriorly, and with a funnel- 
like pit beneath. Umbonal-ventral sulcus 
moderately impressed, sculptured with 
growth lines, and bounded posteriorly by 
a pronounced, shaip ridge. Posterior slope 
rather low and sculptured with well- 
marked growth lines. 



" Named for Frederick M. Bayer, Rosenstiel School 
of Marine and Atmospheric Sciences, University of 
Miami, Miami, Florida, who kindly loaned material 
collected by the research vessels Pillsbunj and Gerda. 



XylophaGAINAE • Turner 



261 



Table 18. Measurements of Xylophaga bayeri. 


Length 


Height 






{mm) 


(mm) 




Location 


8.0 


7.5 


holotyj: 


)e 


4.2 


4.0 


off Mona Island, Puerto Rico 


5.4 


4.9 


Gercla, 


station 266 


6.0 


5.2 


Gerda. 


station 266 


6.0 


5.8 


Gerda, 


station 266 


6.2 


6.0 


Gerda, 


station 266 


6.8 


6.0 


Gerda, 


station 266 


8.0 


S.2 


Gerda, 


station 266 



Inner surface of valves smooth and glis- 
tening. Umbonal— ventral ridge very prom- 
inent and distinctly segmented but not 
greatly enlarged ventrally at the condyle. 
Chondrophore and internal ligament well 
developed. Posterior adductor muscle scar 
elongate oval in outline, tapering dorsally, 
and smooth. Pedal retractor scar broadly 
oval and contiguous with the anterior mar- 
gin of the posterior adductor. Ventral ad- 
ductor scar large, located adjacent to the 
groove margining the umbonal— ventral 
ridge posteriorly. Siphonal retractor scar 
lightly impressed and located just posterior 
to the umbonal— ventral ridge on a level 
with the ventral margin of the beak. An- 
terior adductor scar well marked and cov- 
ering most of the umbonal reflection. 

Mesoplax in young specimens com- 
posed of a large ventral plate with a small, 
triangular dorsal portion (Plate 31, Fig. 5). 
Dorsal plate of the adult is much wider 
than long, with a sharp perphery (Plate 31, 
Figs. 2, 3) and pronounced concentric 
sculpture. 

Siphons capable of complete retraction 
within the valves and of extension proba- 
bly not more than the length of the valves. 
Excurrent siphon truncated, about one 
third length of the incurrent siphon. Lap- 
pets extending from the truncation along 
the dorsal surface of the incurrent siphon, 
finely fringed. Periostracal sheath, if pres- 
ent, extremely thin and lacking calcareous 
or glasslike inclusions. 

Measurements. See Table 18. 

Remarks. Xylophaga bayeri Turner n. 
sp. is a distinctive species apparently not 



closely related to any other species in this 
genus. Its smooth muscle scars, ear-shaped 
mesoplax, truncated excurrent siphon, and 
fringed lappets place it in Group 6 and the 
lack of granular inclusions in the perios- 
tracal sheath of the siphons places it more 
closely to X. globosa and X dorsalis than 
the others in the group. The mesoplax of 
X. bayeri is basically like that of X. dorsalis 
but is greatly extended laterally. 

Only two specimens were obtained from 
the panels placed off Fort Lauderdale, 
Florida, and unfortunately these were not 
sufficiently well preserved for detailed an- 
atomical work. The 14 specimens taken 
from wood dredged off Fowey Rocks were 
in situ, but all were dead and the soft parts 
had disintegrated, although the mesoplax 
was in place. All had typical valves but 
some had unusually wide dorsal plates 
(Plate 31, Figs. 2, 3). Four complete spec- 
imens and about 25 dead and disarticulat- 
ed specimens were extracted from the 
wood dredged off Mona Island. These 
were remarkably uniform in the characters 
of the shell and dorsal plates. 

Range. From off the coast of Florida 
probably throughout the Caribbean, in 
depths ranging from 150 to 365 m. 

Specimens Examined. UNITED STATES, FLOR- 
IDA: 3.2 miles off Fort Lauderdale (26°04'N, 
80°04'W) in 152.4 m (500 ft) in test panel; Ge7'da, 
station 266, off Fowey Rocks (25°39'N, 79°58'W) in 
about 340 ni. PUERTO RICO: Johnson-Smithsonian 
Expedition, station 37, midway between Mona and 
Desecho islands (18°11'55"N, 67°42'50'W) in about 
365 m. 

Xylophaga japonica Taki and Habe 
Plate 32 

Xylophaga japonica Taki and Habe, 1950, Illustrated 
Catalogue of Japanese Shells, No. 7, p. 45, text- 
figs. 6, 7 (Tosa Bay, Shikoku, Japan, in about 100 
fathoms). Holotype, T. Habe Collection; paratype, 
MCZ 194822. 

Distinctive Characters. Posterior slope 
elongate in adult specimens. Umbonal— 
ventral sulcus well impressed, bounded by 
an inconspicuous ridge anteriorly and a 
high narrow ridge posteriorly. Posterior 
adductor muscle scar smooth. Mesoplax 



262 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 19. Measurements of 
Xylophaga japonic a. 



Lengtli Height 
(mm) (mm) 



13.2 10.8 holotype (from Tald and 
Raise, 1950) 
5.8 5.0 paratype Tosa Bay, Shikoku, 

Japan 
3.8 3.0 off Clara Island, South Burma 
2.0 1.8 off Java 



large, covering the upper one half of the 
anterior area, extending over the umbos, 
ear-shaped, moderately inflated, with a 
narrowly rounded peripheiy and distinct 
concentric sculpture (Plate 32, Figs. 2-5). 
Siphons capable of extending two to three 
times the length of the shell. Excurrent si- 
phon about one third the length of the in- 
current, lappets extending along the dorsal 
surface of the incurrent siphon finely 
fringed, opaque white granules imbedded 
along the sides of the siphons. Chimney of 
coarse fecal pellets lining the burrow. 

Measurements. See Table 19. 

Remarks. At the time Taki and Habe 
(1950) described this species, they did not 
have the soft parts and therefore could not 
describe the siphons. However, it is clear 
from their description and figures that the 
species belongs to the dorsalis group 
(Group 6). Taki and Habe related their 
species to X indica Smith, but noted its 
more elongate posterior area. It further 
differs from X indica in its broad meso- 
plax for, according to Knudsen (1961), the 
mesoplax of X. indica is "oblong pear- 
shaped" and his figure [of it] shows it to 
be nearly three times as long as wide. Xy- 
lophaga indica is probably more closely re- 
lated to X. guineensis and X. depalmai n. 
sp., but because the siphons are unknown, 
this cannot be stated definitely. 

Xylophaga japonica appears to be most 
closely related to X. mexicana Dall but dif- 
fers in having a larger, more highly sculp- 
tured mesoplax that extends over the um- 
bos; in having the posterior slope more 
elongate; and in having the ridge bounding 



the umbonal— ventral sulcus inore pro- 
nounced. 

Unfortunately, the material from off 
Java and South Burma noted in the re- 
cords below was poorly presei'ved, but it 
was possible to ascertain the basic char- 
acters of the siphons. The specimens from 
Java had been kept in alcohol since they 
were collected by Mortensen in 1929. The 
specimens from South Burma, dredged by 
the Anton Bniun and taken from a small 
piece of wood and the husk of a nut, had 
been allowed to dry. 

Range. From Tosa Bay, Japan, south and 
west to Burma and Java, in depths ranging 
from 183 to 384 m." 

Specimens Examined. JAPAN: Tosa Bay, Shikoku, p 
in 183 m. JAVA: Danish Java-South African Expedi- 
tion, station 10, off SE tip of Java (08°36'S, 114°34'E) 
in 300 m. BURMA: Anton Bniun, station 23, about 
77 miles W of Clara Island, South Burma (10°30'N, 
96°35'E) in 3S4 m. I 

GENUS XYLOPHOLAS TURNER 1972 

XylopJiolas Turner, 1972, Basteria, 36(2): 97-99. 

Tijpe Species. Xylopholas altenai Turner, 
original designation. I 

Distinctive Characters. Valves and me- f 
soplax typical for the genus Xylophaga. \ 
Animal long, not capable of retraction 
within the valves, with a periostracal si- 
phonal sheath posterior to the valves and | 
a pair of lateral, chitonlike, siphonal plates 
at the posterior end. Siphons short, ex- i 
tending between the plates, the siphonal 
retractor muscles inserted on their inner | 
surface. Gills and visceral mass contained 
between the valves as in typical Xylophaga. j 
Wood-storing cecum large. i 

Remarks. This genus differs from Xylo- 
phaga in having extended excurrent and 
incurrent canals contained in a common j 
sheath with siphonal plates at the posterior 
extremity. These plates probably arose in- 
dependently but may be homologous with i 
the siphonoplax of other pholads (i.e., Pho- j 
ladidea), which was carried posteriorly as i 
the animal elongated. However, the si- I 
phonal retractor muscles of these species 
insert on the valves rather than on the si- 



Xylophagainae • Turner 



263 



phonoplax. The siphonal plates of Xylo- 
pholas probably function as do the pallets 
of the Teredinidae and the siphonoplax of 
other pholadids to close the end of the 
burrow. Embiyological studies are needed 
to prove the affinities of the plates. 

From Xijloredo, Xijlopholas differs in 
having siphonal plates, a periostracal 
sheath on the animal posterior to the 
valves, and in not lining the burrow with 
a chitonlike or calcareous tube. 

Range. To date the genus is known only 
from the type species found off the Lower 
Florida Keys in the western Atlantic and 
in the Gulf of Guinea in the eastern At- 
lantic in depths from 239 to 366 m. (Two 
specimens were dredged in 2,550 m but 
these may have been advectitious.) 

Xylopholas a/tena/ Turner 1972 
Plates 33, 34 

Xylopholas altouii Turner, 1972, Basteria, 36(2): 99- 
103, figs. 1-12 (Gerda, station 66, about 13 miles 
SE of Fowey Rocks, Florida [25°25'N, 79°59'] in 
366 m). Holotype, MCZ 279315. 

Distinctive Characters. Aniinal elon- 
gate, not capable of retraction within the 
valves, and with lateral, paddlelike siphon- 
al plates at the posterior end. Shell similar 
to that of Xijlopliaga. Mesoplax composed 
of two flat, elongate plates that are held in 
place by the periostracum extending be- 
tween the beaks dorsally. Posterior adduc- 
tor muscle scar large and with transverse 
forking impressions. Young carried on the 
ventral surface just posterior to the valves. 

Description. Shell globose, reaching 2.5 
mm in length, thin, fragile, and with a rel- 
atively heavy, golden-brown periostracum 
covering the valves and mesoplax. Pedal 
gape angle about 90°. Beaked portion of 
the anterior slope recurved dorsally and 
sculptured with numerous strong, dentic- 
ulated ridges. Umbonal reflection narrow. 
Umbonal— ventral sulcus narrow and only 
slightly impressed. Disc and posterior 
slope sculptured with fine growth lines 
only. 

Inner surface of valves smooth and glis- 
tening. Umbonal-ventral ridge low, indis- 



tinct except near the ventral margin, 
slightly segmented, and with a small ven- 
tral condyle. Chondrophore and internal 
ligament well developed. Posterior adduc- 
tor muscle scar large, covering most of the 
posterior slope, elongate oval in outline 
and marked with transverse, forking im- 
pressions. Pedal retractor scar irregularly 
and broadly oval and located about mid- 
way on the anterior margin of the poste- 
rior adductor scar. Siphonal retractor mus- 
cles inserted on the siphonal plates and 
collar. 

Mesoplax small, not filling the gape be- 
tween the beaks, composed of two flat, 
elongate, subrectangular plates, somewhat 
pointed posteriorly, sculptured with fine 
transverse ridges, covering the dorsal sur- 
face of the anterior adductor muscle and 
held in place by the periostracum. 

Animal long, with a periostracal sheath 
covering the portion posterior to the valves 
and with a pair of lateral, paddle-shaped, 
chitonlike plates at the posterior end 
(Plate 33, Fig. 4). The siphons extend be- 
tween the plates, and the siphonal retrac- 
tor muscles insert on the inner surface of 
them. Siphons separate, excurrent siphon 
longer than the incurrent siphon, the ap- 
ertures of both with fine cirri. 

Gills and visceral mass contained entire- 
ly between the valves, the portion of the 
animal extending beyond the valves com- 
posed of a dorsal excurrent and a ventral 
incurrent canal combined in a common 
muscular and periostracal sheath, with a 
chitinous collar and two lateral, paddle- 
shaped plates posteriorly. 

Measurements. See Table 20. 

Remarks. Isolated valves of this species 
would be difficult if not impossible to dis- 
tinguish from several species of Xylopha- 
ga; however, its reduced, flat mesoplax, 
elongate soft parts, and siphonal plates 
readily distinguish it from all other species 
in the Xylophagainae. Nothing is known of 
the biology of the species except that it has 
a large wood-storing cecum and, therefore, 
probably utilizes wood as food. The young 
are held within the burrow to the late ve- 



264 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 20. Measurements of 
Xylopholas altenai. 



Length 

(mm) 



Height 



1.8 


1.8 


holotype 


1.0 


1.0 


Atlantique Sud, station 147 


1.5 


1.4 


Galathea, station 52 


1.2 


1.1 


Gerda, station 66 


1.9 


1.8 


Gerda, station 266 


2.0 


2.1 


Gerda, station 266 


2.0 


2.5 


Gerda, station 266 


2.5 


2.5 


Gerda, station 266 



liger stage when the foot has developed. 
They are attached to the ventral surface of 
the animal just posterior to the valves. The 
number of attached young ranged from 
two to eight and they averaged 0.28 mm 
in length. The hinge plate of the young is 
well developed, with two teeth and a sock- 
et in the right valve and a corresponding 
single tooth and two sockets in the left 
valve (Plate 33, Figs. 1, 2). 

About 60 specimens were taken at the 
three stations off Florida and 10 speci- 
mens were taken from the three stations 
in the Gulf of Guinea. The shells of many 
of the specimens were in poor condition 
and the valves had largely dissolved, pos- 
sibly a result of the wood being veiy acid. 
However, the characteristic siphonal plates 
readily identified the species. The two 
specimens taken from a coconut husk 
dredged off Sao Tome in 2,550 m may 
have been advectitious; this species may 
not typically occur at that depth. Neither 
specimen carried young. The shelf in this 
area is very narrow and steep so that plant 
material could easily be carried into deep 
water. All other records are from depths of 
239-366 m. 

Range. Known only from off the Florida 
Keys in the western Atlantic and the Gulf 
of Guinea in the eastern Atlantic in depths 
ranging from 239 to 2,550 m. 

Specimens Examined. Western Atlantic: UNITED 
STATES, FLORIDA: Gerda, station 266, off Fowey 
Rocks, about 16 miles SE of Miami (25°39'N, 
79°58'W) in 340 m; Gerda, station 66, off Turtle 
Reef, about 13 miles SE of Fowey Rocks (25°25'N, 



79°59'W) in 366 ni; Gerda, station 220, about 30 
miles S of Alligator Reef (24°25'N, 80°33.5'W) in 311 
m. Eastern Atlantic: GABON: Galathea, station 52, 
off Port Victoria, Sao Tome Island (1°42'N, 7°51'E) 
in 2,550 m (in coconut shell); Atlantique Sud, station 
147, about 45 miles N Port Gentil, Cape Lopez (0°S, 
8°58'E) in 250 m; Atlantique Sud, station 154, about 
35 miles NE of Port Gentil (0°15'S, 8°47'E) in 239 m. 

GENUS XYLOREDO TURNER 1972 

Xyloredo Turner, 1972, Breviora, 397: 3 (type species, 
Xyloredo nooi Turner, original designation); Turner, 
1973, Science, 180: 1377-1379. 

Distinctive Characters. Species in this 
genus are characterized by having typical 
Xylophaga shells, which lack apophyses 
and have a mesoplax composed of two flat 
plates, and by making a long teredolike 
burrow. The posterior two thirds of the 
burrow has a thin calcareous lining, 
marked with distinct growth lines and cov- 
ered with periostracum that extends to the 
calcareous portion anterior as a band. The 
part of the animal extending beyond the 
valves into the calcareous tube is covered 
by a golden-brown periostracal sheath that 
is continuous anteriorly with the covering 
of the valves and posteriorly with the per- 
iostracum of the tube. A fold of the mantle 
is attached to the growing end of the tube 
where both periostracum and calcium are 
added. In young specimens the tube may 
be composed entirely of periostracum. 

Posterior to the valves, the combined in- 
current and excurrent canals extend the 
length of the tube and are attached lightly 
to it at the base of the short separate si- 
phons. Two dorsolateral ridges within the 
incurrent canal appear to be ciliated and 
possibly aid in water transport. 

Remarks. Members of this genus differ 
from Xylophaga and Xylopholas in making 
a burrow that may reach more than 30 
times the length of the shell and is lined 
with a calcareous tube. Several species of j 
Xylophaga make a burrow more than five 1 
times the length of the shells and form a | 
chimney composed of mucous-cemented [ 
fecal pellets at the posterior end of the | 
burrows. These are not homologous with | 
the calcareous tubes of Xyloredo but rath 



Xylophagainae • Turner 265 



er with the chimney of rock-boring pho- 
lads, as in Parapholas Conrad (Turner, 
1955: 123). 

The discovery of this teredohke genus 
in the Pholadidae requires a reexamination 
of the fossil teredinids, especially those re- 
corded as having ringed tubes. On the ba- 
sis of our present knowledge, it may be 
impossible to distinguish Xyloredo from 
teredinids in fossilized wood. However, if 
tubes are present, a microscopic analysis 
of their structure may aid in distinguishing 
between them because teredinid tubes are 
amorphous, whereas tubes of Xyloredo 
have a definite structure with growth rings 
and periostracum. Certainly Xyloredo 
should be considered when examining 
drilled wood thought to have come from a 
deep water fossilized deposit. 

Although Xyloredo superficially resem- 
ble the Teredinidae, this is entirely con- 
vergent and does not in any way indicate 
relationship, nor does it suggest the evo- 
lution of the Teredinidae from the Xylo- 
phagainae. The latter lack apophyses and 
pallets, and have a mesoplax. In addition, 
none of the visceral mass or gills of the 
Xylophagainae extend beyond the valves 
posteriorly as they do in the Teredinidae. 

Range. To date three species of Xylo- 
redo are known, two in the western Atlan- 
tic and the other in the eastern Pacific. All 
occur at depths greater than 1,500 m. 

Xyloredo noo/ Turner 
Plate 35 

Xyloredo nooi Turner, 1972, Breviora, 397: 5—7, pis. 
1, 2 (Tongue of the Ocean, about 4 miles off north- 
east tip of Andros Island, Bahama Islands [25°.54'N, 
77°49'W] in 1,737 m). Holotype, MCZ 279631; 
paratypes from the same and other panels exposed 
at the same locality, MCZ 279632, 279633, 279634, 
and 279635, and the Zoological Museum, Univer- 
sity of Copenhagen; Turner, 1973, Science, 180: 
1377-1379. 

Distinctive Characters. Burrow long, 
teredinidlike, lined with a thin calcareous 
tube marked with growth rings and cov- 
ered with periostracum. Shell similar to 
Xylophaga, anterior slope narrow, umbo- 
nal— ventral sulcus lightly impressed, pos- 



terior slope high and reflected dorsally 
Posterior adductor muscle scar subellipti- 
cal, set high on the posterior slope, and 
divided into two distinct areas. Disc sep- 
arated from the posterior slope by a 
groove on the inner surface of the valves. 
Mesoplax small, the two flat triangular 
plates composed almost entirely of perios- 
tracuiTi. Periostracal sheath between the 
valves and the tube smooth. 

Description. Shell globose, valves reach- 
ing 10 mm in length and 10.5 mm in 
height, thin, fragile; umbos prominent. 
Periostracum relatively thick, golden- 
brown, glistening, and covering entire 
valve. Pedal gape angle about 110°. Ante- 
rior slope sculptured with numerous den- 
ticulated ridges, there being 24 on the ho- 
lotype. UiTibonal— ventral sulcus narrow, 
slightly impressed, and sculptured with 
fine, irregular growth lines. Posterior slope 
high, reflected near the dorsal margin, and 
sculptured with fine growth lines. Umbo- 
nal reflection rather wide, thin, adhering 
to the valves in the umbonal area, free an- 
teriorly. 

Inner surface of valves smooth and glis- 
tening. Umbonal— ventral ridge narrow, 
high, and segmented. Chondrophore and 
internal ligament prominent. Disc separat- 
ed from the posterior slope by a pro- 
nounced narrow groove extending from 
the umbo to the posterior ventral margin. 
Posterior adductor muscle scar large, ellip- 
tical, and divided into two areas, the upper 
part marked with irregular impressions, 
the lower with regular chevron-shaped im- 
pressions. Anterior adductor scar covering 
most of the umbonal reflection. Pedal re- 
tractor scars elongate, the ixiuscles insert- 
ing in the groove separating the disc from 
the posterior slope. 

Mesoplax small, flat, the two broadly tri- 
angular plates composed almost entirely of 
periostracum and located anterior to the 
umbos. 

Burrow long, teredinidlike, and lined 
with a thin calcareous tube that is sculp- 
tured with distinct growth rings and cov- 
ered with periostracum that extends as a 



266 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 21. Measurements of Xyloredonooi. 



Length 


Hei2;ht 




(mm) 


(mm) 


Location 


5.0 


5.1 


paratype 


6.7 


6.9 


paratype 


7.2 


7.8 


paratype 


9.5 


9.8 


paratype 


9.5 


10.0 


holotype 



border at the anterior end. Tube in veiy 
young specimens composed entirely of 
periostracum. Between the valves and the 
anterior end of the tube the animal is cov- 
ered with a smooth periostracal sheath, 
which is continuous anteriorly with the 
covering of the valves and posteriorly with 
the tube. Siphons short, separate, and ap- 
parently with a few small cirri. 

Measurements. See Table 21. 

Remarks. Xyloredo nooi is related to 
both Xyloredo naceli Turner from the east- 
ern Pacific and X ingolfia Turner from the 
North Atlantic. Xyloredo nooi differs from 
them in having a much thinner burrow lin- 
ing, a high, reflected posterior slope on the 
valves, and a proportionately smaller, more 
highly placed and divided posterior adduc- 
tor muscle scar. In addition, the periostra- 
cal sheath extending between the valves 
and the calcarious tube is smooth. 

The larvae of X. nooi apparently do not 
rise very high in the water column, inas- 
much as a panel 25 ft off the bottom 
showed no trace of them, whereas the 
panel directly beneath it on the bottom 
was riddled. On the basis of the prodis- 
soconch still visible on some of the speci- 
mens, it would appear that the lai'vae are 
similar to those ofXylophaga. No evidence 
was found in the three riddled panels ex- 
amined that the young were brooded or 
held in the tubes. 

The burrows resemble those of teredi- 
nids and intertwine with each other but 
basically follow the grain of the wood. 
They may reach 200 mm in length and 15 
mm in diameter at the anterior end. The 
largest tube with intact shells and animal 
remaining was 145 mm long; the valves 



were 9.5 mm in length and 10 mm in 
height. The panels in the Tongue of the 
Ocean were submerged for 34 months and 
inasmuch as the larger burrows were emp- 
ty or contained only fragments of shells, 
the length of life may be about 2—2.5 
years. The calcareous lining of a burrow 
122 mm long extends to about 30 nam 
from the anterior end, thus leaving room 
for boring activities and changing the di- 
rection of the burrow. 

Range. Knowii only from the type lo- 
cality. 

Specimens Examined. BAHAMA ISLANDS: 
Tongue of the Ocean, about 4 miles off NE tip of 
Andros Island (24°54'N, 77°49'W) in 1,737 m. 

Xyloredo ingolfia Turner 
Plates 36, 37 

Xyloredo ingolfia Turner, 1972, Breviora, 397: 7-9, 
pis. 3-5 (from wood dredged by the Ingolf Expe- 
dition at station 67, south of Eyrabakld, Iceland 
[61°30'N, 22°30'W] in 975 fathoms [1,783 m]). Ho- 
lotype, MCZ 279636; paratypes, MCZ 279637, and 
the Zoological Museum, University of Copenha- 
gen; Turner, 1973 Science 180: 1377-1379. 

Description. Shell globose, valves reach- 
ing 2.5 mm in length and 2.0 mm in 
height, thin, fragile, with prominent um- 
bos; thin, glistening, almost colorless per- 
iostracum covering disc and posterior 
slope. Beaked portion of anterior slope 
wide, extending more than one half dis- 
tance to ventral margin; sculptured with i 
close-set and very finely denticulated ridg- 
es. Posterior portion of anterior slope 
about two thirds width of beak, sculptured ! 
with close-set ridges that extend to very 
slightly impressed umbonal— ventral sulcus. 
Disc sculptured with well-marked growth 
lines. Posterior slope small, low, and not 
clearly demarcated on outer surface of 
valve. Umbonal reflection thick, narrow, 
short, and free except at posterior end. 

Inner surface of valves smooth, slightly 
shiny to chalky (perhaps owing to long ' 
preservation). Umbonal-ventral ridge wide, 
flattened, often varying in width, irregularly 
segmented, and not enlarged at ventral 
condyle. Chondrophore and internal liga- 



Xyloph AGAIN AE • Turner 



267 



Table 22. Measurements of 
Xyloredo ingolfia. 



Range. Known only from the type loeal- 



Lrimtll 


Heidit 




(mill) 


(mm) 


Location 


2.5 


2.3 


holotype 


2.5 


2.0 


paratype 


2.1 


2.0 


paratype 


2.0 


1.9 


paratype 


1.5 


1.4 


paratype 


1.5 


1.2 


paratype 



ment large. Disc not clearly separated from 
posterior slope. Posterior adductor muscle 
scar large, slightly raised, elliptical, extend- 
ing nearly to ventral margin, with irregular, 
transverse impressions. Anterior adductor 
muscle scar covering umbonal reflection. 
Siphonal retractor muscle scars not im- 
pressed. Pedal retractor muscle scar small, 
elongate to oval, and located just anterior 
to posterior adductor muscle scar. Meso- 
plax of two veiy small, narrow, subrectan- 
gular, flat, calcified plates lying on dorsal 
surface of anterior adductor muscle. 

Burrow 10—15 times length of valves; cal- 
careous tubular lining three fourths length 
of bruTow. Tube relatively heavy, marked 
with rmiform, close-set, raised rings, and 
covered with light tan periostracum that ex- 
tends anteriorly as border. Portion of ani- 
mal between valves and tube covered by 
tliin, irregrflarly ridged periostracal sheath. 
Siphons short; incurrent siphon slightly lon- 
ger than excurrent siphon. Protoconch 
large, medium golden-brown, and sculp- 
tured with fine, concentric ridges. 

Measurements. See Table 22. 

Remarks. This species differes from Xy- 
loredo naceli in having a less well-devel- 
oped posterior slope, a shallow, indistinct 
umbonal— ventral groove, a flattened um- 
bonal— ventral ridge, and in having the 
valves longer than high. It differs from X. 
nooi in having valves longer than high, in 
having a low, rounded posterior slope, in 
lacking the distinct groove on the inner sur- 
face separating the disc from the posterior 
slope, and in having the plates of the me- 
soplax sub rectangular and well calcified. 



ity. 



Specimens Examined. ICELAND: Ingolf Expedi- 
tion, station 67, S of Eyrabakld (61°30'N, 22°30'W) 
in 1,783 m. 

Xyloredo nace// Turner 
Plate 38 

Xyloredo naceli Turner, 1972, Breviora, 397: 9—11, pi. 
6, figs. 1-5 (USNCEL STU 1-4 about 30 miles S 
of San Miguel Island, off Port Hueneme, Santa 
Barbara Islands, California [33°46'N, 120°45'W] in 
6,800 ft [2,072.6 m] from panels submerged from 
June 1964 to July 1965). Holotype, MCZ 279638; 
paratope, 279639. 

Distinctive Characters. Burrow teredi- 
nidlike, lined with a thin calcereous tube 
marked with growth rings and covered 
with periostracum. Shell similar to Xijlo- 
phaga, with a narrow, slightly impressed 
umbonal— ventral sulcus. Posterior adduc- 
tor muscle scar elliptical, almost complete- 
ly covering the posterior slope and uni- 
formly marked with transverse impres- 
sions. Mesoplax small, the two flat, trian- 
gular plates composed largely of 
periostracimi. Periostracal sheath covering 
the animal between the valves and the 
tube papillose. 

Description. Shell globose, reaching 1.5 
mm in length, thin, fragile, white, with a 
thin, pale yellow periostracum. Pedal gape 
angle about 100°. Anterior slope sculp- 
tured with 8—12 widely spaced, pro- 
nounced, denticulated ridges. Umbonal— 
ventral sulcus narrow and only slightly im- 
pressed. Discs and posterior slope sculp- 
tured with fine growth lines only. 

Inner surface of valves smooth and glis- 
tening. UiTibonal— ventral ridge narrow, 
high, and indistinctly segmented. Chon- 
drophore and internal ligament well de- 
veloped. Posterior adductor muscle scar 
elliptical, extending from the dorsal nearly 
to the ventral margin of the posterior slope 
and regularly marked with transverse im- 
pressions. Pedal retractor scar not im- 
pressed. Ventral adductor only lightly im- 
pressed and located just posterior to the 



268 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 



Table 23. Measurements of Xyloredo naceli. 



Length 

(mm) 



1.1 

1.0 

1.2 
1.5 


1.2 
1.0 
1.3 
1..5 


holotype 
paratype 
paratype 
paratype 



umbonal— ventral ridge. Siphonal retractor 
muscles inserting on the tube. 

Mesoplax small, thin, set between and 
anterior to the umbos, the two triangular 
plates composed almost entirely of perios- 
tracum. Burrow of largest specimen about 
six times the length of the shell. Calcare- 
ous tube lining the burrow thin, distinctly 
marked with growth rings, and covered 
with periostracum that extends as a border 
anteriorly. Aperture of the burrow small, 
round, the white lining visible within. The 
portion of the animal between the valves 
and the calcareous tube covered with a pa- 
pillose periostracal sheath that is continous 
with the periostracum covering the valves 
anteriorly and the periostracal border of 
the tube posteriorly. Siphons short, of 
equal length, and apparently lacking cirri. 

Measurements. See Table 23. 

Remarks. This species is most closely re- 
lated to X nooi Turner from the western 
Atlantic. Only eight specimens of X. naceli 
were found and they were all very small, 
but these appear to be sufficiently distinct 
to consider them members of a separate 
species. Xyloredo naceli differs from X 
nooi in having a heavier tube that is cal- 
careous, even in very yotmg specimens; 
and a posterior adductor muscle scar that 
is proportionately larger and not divided 
into two areas. In addition, it lacks the pro- 
nounced internal groove separating the 
disc from the posterior slope, and the per- 
iostracal sheath covering the animal ante- 
rior to the tube is papillose. 

Nothing is known of the biology of the 
species except that at the sites where they 
were collected the temperature of the wa- 
ter was 2.1° C, the salinity was 34.52%o, 
the dissolved oxygen content was 1.26 



ml/L, the pH was 7.84, and the hydrostatic 
pressure 3000 psi (Mru-aoka, 1966b). 

The embryonic valves still visible on 
some of the specimens suggest that the 
mature larvae are similar to those in Xy- 
lophaga. Xylophaga niuraokai was the 
most common borer in the panels from 
which the X. naceli were taken. 

Range. Known only from the type lo- 
cality. 

Specimens Examined. UNITED STATES, CALI- 
FORNIA: USNCEL STU 1-4 S of San Miguel Island, 
Santa Barbara Islands (33°46'N, 120°45'W) in 
2,072 m. 

ACKNOWLEDGMENT 

This paper was prepared with the aid of 
funds received from the Department of 
the Navy, Biology Branch, Office of Naval 
Research, ONR grant N00014-91-J-1402, 
Biological Studies on Marine Boring and 
Fouling Mollusks. 

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Xylophagainae • Turner 269 



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270 Bulletin Museum of Comparative Zoologt/, Vol. 157, No. 4 




2 mm 
Plate 1 . Xylophaga concava Knudsen from Pillsbury, station 526. 



1 mm 



Figure 1 . Dorsal view of apposed valves showing the concave posterior slope and erect mesoplax. Figure 2. Posterior end of 
valve and siphons. Figure 3. Enlargement of the siphonal openings to show the six large cirri on the excurrent siphon and the 
double row of small cirri on the incurrent siphon. 



Xylophagainae • Turner 271 




Plate 2. Xylophaga gerda Turner n. sp. from Gerda, station 499. 

Figure 1 . Lateral view of holotype showing the attachment of the posterior adductor muscle through the thin valve, the mesoplax 
that does not extend above the umbos, and the fecal cylinder in the excurrent canal. Figure 2. Dorsal view of the holotype 
showing the mesoplax. Figure 3. Enlargement of the posterior end of the siphons. Figure 4. Diagrammatic cross-section 
through the siphons and the fecal cylinder. Figure 5. Three-quarter view of holotype showing the inflated umbos and the simple 
curved plates of the mesoplax. Figure 6. A relatively smooth chimney composed largely of periostracum, with a thin coating of 
fecal material. Figure 7. A thick chimney, built in sections with "leaves" of periostracum extending at the anterior end of each 
section. 



272 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4 




1 mm 
Plate 3. Xylophaga gerda Turner n. sp. from Pillsbury, station 944. 



0.5 mm 



Figure 1 . Outer view of right valve. Figure 2. Inner view of left valve, showing posterior adductor muscle scar