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US ISSN 0006-9698

CAMBRIDGE, MAss.

11 NovEMBER 2011

NUMBER 528

' SHEAROGOVEA, A NEW GENUS OF CYPHOPHTHALMI (ARACHNIDA,
OPILIONES) OF UNCERTAIN POSITION FROM OAXACAN CAVES, MEXICO

GONZALO GIRIBET!

ABSTRACT.

Shearogovea gen. nov. is erected for Neogovea mexasca Shear, 1977, a troglobitic cyphophthalmid

species from a cave system in Oaxaca, Mexico. The new genus does not show affinity to Neogovea Hinton, 1938, as it
lacks the characteristic toothed claw of leg II or the fusion of the coxae of legs II to those of legs III (which are in turn
fused to coxae of legs IV). Shearogovea gen. nov. is probably not related to other Neotropical neogoveid genera, but

its exact phylogenetic position remains unresolved.

KEY WORDS:

INTRODUCTION

Shear (1977) reviewed the Neotropical
genus Neogovea Hinton, 1938 and described
two new species, N. kamakusa Shear, 1977
from Guyana, and N. mexasca Shear, 1977
from a cave in Oaxaca, Mexico. Additional
information on the latter species, originally
described for two female individuals, was
provided in a later paper based on a larger
collection of specimens, including males,
from a nearby cave (Shear, 1980). In the
same paper, a new familial system was

"Museum of Comparative Zoology, Department of
Organismic and Evolutionary Biology, Harvard Univer-
sity, 26 Oxford Street, Cambridge, Massachusetts 02138,
U.S.A.; e-mail: ggiribet@oeb.harvard.edu.

troglobite; Mexico; new genus; Neogoveidae; Sironidae

established for Cyphophthalmi, and Neogo-
veidae Shear, 1980 was designated for the
genera Metagovea Rosas Costa, 1950, Paro-
govia Hansen, 1921 [misspelled as Paragovia]
and Neogovea, including N. mexasca, despite
acknowledging that it lacks characters used
to diagnose the neogoveid genera (Shear,
WS).

The Mexican species differs from its
supposed South American counterparts in
several characters, also shared with the
African Parogovia, later used to rediagnose
Neogoveidae (Giribet, 2007b). Thus UN.
mexasca was considered of uncertain phylo-
genetic position in several studies (Giribet
and Boyer, 2002; Schwendinger et al., 2004;
Giribet and Kury, 2007; DaSilva et al., 2010)

© The President and Fellows of Harvard College 2011.

2 BREVIORA

esi

Bes

Figure 1. Shearogovea mexasca (Shear, 1977), fe-
male holotype, American Museum of Natural History.
A, dorsal view; B, ventral view.

and recently excluded from Neogoveidae
(Benavides and Giribet, 2007). However,
some of these characters were considered
adaptations to troglobite lifestyle, and was
thus considered to be an aberrant neogoveid
(Shear, 1979). Here, Shearogovea gen. nov. is
erected for N. mexasca Shear, 1977 on the
basis of detailed morphological analysis
using scanning electron microscopy (SEM),
but its familial assignment remains uncertain
given the currently available data.

MATERIALS AND METHODS

The material used in this study belongs to
the collection of the American Museum of
Natural History and the Museum of Com-
parative Zoology. The holotype specimen

No. 528

feo BOe A TEAS Goa
100mm

a) Vee (aed od ek fit oe ed ol
1.00mm

Figure 2. Shearogovea cf. mexasca from Cueva de
la Finca. A, male, ventral view; B, female, ventral view.

was photographed using a JVC KY- F70B
digital camera mounted on a Leica MZ 12.5
stereomicroscope. A series of images (about
10) were taken at different focal planes and
assembled with the dedicated software pack-
age Auto-Montage Pro Version 5.00.0271
(Syncroscopy, Frederick, MD, USA). Scan-
ning electron micrographs were taken with a
Hitachi $4700 FE-SEM, after sputter coat-
ing with gold/palladium.

Shearogovea Giribet, gen. nov.
Figures 1-4

Neogovea, Partim. Shear, 1977:172-175,
figs. 16-24; Morales Soto, 1980:36—37;
Shear, 1980:15-17, figs. 21-25; Cokendol-
pher and Lee, 1993:5; Rambla and Juberthie,
1994:217; Kury and Cokendolpher, 2000:
150-151; Gziribet, 2000:71; Giribet and
Boyer, 2002:110-116, 118, 121, 127-128;

2011 SHEAROGOVEA FROM MEXICO 3

Figure 3. Shearogovea cf. mexasca from Cueva de la Finca. A, prosomal sternal complex, male; B, prosomal
sternal complex, female; C, anal region, male; D, spiracle; E, left chelicer, male; F, detail of cheliceral fingers, male;
G, left palp, male.

4 BREVIORA

Giribet,
2011:49.

‘Neogovea’. Schwendinger et al., 2004:
1422: Benavides and Giribet, 2007:2.

““Neogovea’’. Giribet and Boyer, 2002:110—
116, 118, 121, 127, 128; Giribet and Kury,
2007:68, 83; DaSilva et al., 2010:52.

Material Studied. The type material of N.
mexasca consists of a female holotype and a
female paratype from Cueva del Nacimiento
del Rio San Antonio, 10 km SSW of Acatlan
de Pérez Figueroa, Oaxaca, Mexico, col-
lected 31 December 1973 by James Reddell,
William Elliott, and Roy Jameson, deposited
in the American Museum of Natural History
(Shear, 1977). Another collection from
Cueva de la Finca, 10 km SW of Acatlan
de Pérez Figueroa, Oaxaca, Mexico, col-
lected 31 December 1976 by James Reddell,
David McKenzie, and Andy Grubs, also
deposited in the American Museum of
Natural History, includes five males and
nine females. A male and a female mounted
in SEM stubs have been transferred to the
Museum of Comparative Zoology under
accession numbers MCZ 124538 and
124539, respectively. No other collections
_ of this or related species are known.

Diagnosis. Medium-sized cyphophthal-
mids with elongated appendages (Fig. 1),
without eyes or eye lenses; ozophores of type
2 (sensu Juberthie, 1970) (Fig. 1B), entirely
ornamented, with subterminal ozopore. An-
terior margin of carapace concave dorsally,
leaving the base of the chelicerae and the
dorsal crest clearly exposed (Fig. 1A, B).
Transverse prosomal sulcus inconspicuous
(Fig. 1A). Transverse opisthosomal sulci
present. Middorsal, longitudinal opisthoso-
mal sulcus inconspicuous (Fig. 1A).

Coxae of leg II free, not fused to coxae of
legs III (Figs. 2, 3A, B). Sternum absent
(Figs. 2A, 3A). Proximal end of coxae I of
males not meeting along the midline; all
other coxae meeting along the midline; male

2007a:94; Jocqué and Jocque,

No. 528

gonostome semicircular, as large as the
proximal ventral part of coxae of legs IV.
Coxae II-III and IJJ-IV with endites running
along their suture. Anterior projections of
male coxae IV endites on gonostome wall
(Fig. 3A). Spiracles circular, of the closed
type (Fig. 3D). Sternites 8 and 9 and tergite
IX fused into a corona analis (Fig. 3C). Anal
plate and anal region without conspicuous
modifications or changes in ornamentation
pattern. Opisthosomal exocrine glands of
males absent (both sternal or anal glands).
Hansen’s organ absent.

Chelicerae of the protruding type (sensu
Giribet, 2003) (Fig. 1A, B), with one dorsal
and one basal process; with the basal and
second cheliceral segments elongate, almost
of uniform depth; first cheliceral segment
ornamented at the first half, near the base,
and in the ventral portion almost during its
entire length; broadest part of second chelic-
eral segment neither near the base nor the
joint with the cheliceral finger (Fig. 3E);
dentition of mobile digit uniform (Fig. 3F).
Palp trochanter without ventral process; first
and second article of palp ornamented

(Fig. 3G).
Metatarsus of all legs ornamented
(Fig. 4A, D); tarsi of all legs smooth

(Fig. 4A—G), without a distinct solea on leg
I (Fig. 4C); claws of all legs smooth, without
comblike modifications or lateral pegs
(Fig. 4C—G). Tarsus IV of males not divided
(Fig. 4A); Rambla’s organ absent. Adeno-
style conspicuous, thin, of the lamellar type
(Fig. 4B); located in the first quarter of the
tarsal length, but not in a basal position
(Fig. 4A).

Spermatopositor with short ventral plate
and without ventral microtrichiae, lacking
moveable fingers (see Shear, 1980, figs. 24—
25). Ovipositor typical, with terminal sense
organs (Shear, 1977:174—-175).

Included Species. Shearogovea mexasca
(Shear, 1977) is the type and only species in

2011

CRORE RED OS
100um

SHEAROGOVEA FROM MEXICO 5

Ps Peis
um

Figure 4. Shearogovea cf. mexasca from Cueva de la Finca. A, left leg 1V, male; B, adenostyle; C, left leg I, male;
D, left leg IV, female; E, left claw II, male; F, left claw III, male; G, left claw IV, male.

the genus (type locality: Cueva del Naci-
miento del Rio San Antonio). Whether the
second collection from Cueva de la Finca, a
nearby cave, represents the same species or
not, remains unsolved, as no males are
known from the type locality. These two
caves are not in the same hill, but the two
hills are separated at the surface by | km of
valley fill comprised of soil and volcanic
boulders, but in the subsurface the limestone
is likely contiguous, as is the case of the
troglobitic tarantula Hemirrhagus reddelli

(Gertsch, 1973) and the troglobitic schizomid
Stenochrus firstmani (Rowland, 1973) (Row-
land and Reddell, 1980). However, until
males from both caves or molecular data

become available, I refer to the specimens

from the second locality as Shearogovea cf.
mexascd.

Etymology. The generic name is a combi-
nation of the last name of arachnologist and
myriapodologist William Shear, to whom
this genus is dedicated—in honor of his
many contributions to the study of Cy-

6 BREVIORA

phophthalmi, which include, among many
other aspects, four genera and 30 species—
and the termination -ogovea, typical of
neogoveid and ogoveid species. Gender is
feminine.

Remarks. The type species of the genus
was originally described in the genus Neogo-
vea Hinton, 1938, although in the original
description Shear (1977:175) stated that
“The general appearance of the animal
suggests intermediacy between Metasiro
americanus and the other species of Neogo-
vea, and a new generic name may be
warranted when males are discovered.” In
the following paper describing the males
from a nearby cave, Shear (1980:17) also
pointed out that “The adenostyle of N.
mexasca is not like that of the epigean
species, but more like that found in species
of Siro. I cannot say if this represents a
specialization or the retention of an ancestral
form. Certainly, N. mexasca is far separated
from the South American species and could
be a very ancient relict of an original
Neogovea stock—predating both the origin
of the brushlike adenostyle and the extensive
_ coxal fusion of the South American species.”

After studying the evolution of Neogovei-
dae and its related families (Ogoveidae and
Troglosironidae) (Giribet et al., in press) we
now understand that major differences exist
between the Amazonian genus Neogovea and
Shearogovea gen. nov. These differences are
not restricted to the adenostyle, so charac-
teristic in the genus Neogovea (see Jocqué
and Jocquey 2011) Ther ince coxac rill
distinguishes Shearogovea from most neogo-
veid genera, although the Brazilian Canga
DaSilva, Pinto-da-Rocha & Giribet, 2010
and the North American Metasiro Juberthie,
1960 also show this condition (Juberthie,
1960; DaSilva et al, 2010). This character
state is shared with pettalids, troglosironids,
and most sironids (except for Iberosiro de
Bivort & Giribet, 2004 and Paramiopsalis

No. 528

Juberthie, 1962), but differs from stylocellids
and ogoveids, which, like most neogoveids,
show fusion to the coxae of legs III (which
are always fused to coxae of legs IV) (Giribet
and Boyer, 2002). The lack of dentition in
the claw of walking leg II and lack of any
sort of opisthosomal exocrine glands, now
interpreted as synapomorphies of the clade
including these three families, also removes
Shearogovea from the family Neogoveidae,
as proposed by Benavides and Giribet
(2007). The lack of jagged dentition in the
cheliceral mobile digit and the absence of
opisthosomal exocrine glands are two other
character states differing from neogoveids
and troglosironids.

The placement of Shearogovea gen. nov. in
the cladistic analysis of Giribet and Boyer
(2002) was rather unresolved, and under
their “root 2”, would have resulted in an
unresolved position with Parasiro and a
clade containing ogoveids, neogoveids, trog-
losironids and stylocellids. A recent analysis
combining molecular data and morphology
(the former not available for Shearogovea
gen. nov.) shows again an unstable phyloge-
netic position of Shearogovea gen. nov., but
it does not group with neogoveids or with
any of the related families Ogoveidae and
Troglosironidae (Giribet et al., in press). A
relationship to the Laurasian family Sironi-
dae is, however, plausible, given the data
available at the moment, which would
reinforce the idea of Shearogovea gen. nov.
being a Laurasian relict.

ACKNOWLEDGMENTS

Abel Pérez-Gonzalez, Rudy Jocqué, and
Ricardo Pinto-da-Rocha provided comments
that helped to improve an earlier version of
this article. Samantha Edelheit and Jonathan
Losos are acknowledged for their help with
the handling and publication of this paper.
This work has been possible in large part due

2011

to a grant from the National Science
Foundation (DEB-0236871).

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