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Revision and cladistic analysis of the genus Hemirrhagus Simon, 1903 (Araneae, Theraphosidae, Theraphosinae)

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Bull. Br. arachnol. Soc. (2003) 12 (8), 365–375
Revision and cladistic analysis of the genus
Hemirrhagus Simon, 1903 (Araneae,
Theraphosidae, Theraphosinae)
Fernando Pérez-Miles
Sección Entomología, Facultad de Ciencias,
Iguá 4225, 11400 Montevideo, Uruguay
and
Arturo Locht
Laboratorio de Acarología, Facultad de Ciencias,
Universidad Nacional Autónoma de Mexico,
Coyoacán 04510, D.F., Mexico
Summary
The genus Hemirrhagus Simon, 1903 is reviewed on the
basis of the study of types and new collections from Mexico.
Six new species are described and illustrated. Hemirrhagus
peruvianus Chamberlin, 1916 and Hemirrhagus major
Chamberlin, 1916, both from Peru, are here removed from
Hemirrhagus because they lack the diagnostic characters of
the genus; they are transferred to Homoeomma Ausserer,
1871 and Hapalotremus Simon, 1903 respectively (new
combinations). As aresult of the study of types of all species
of Spelopelma Gertsch, 1982, found mostly in caves, this
genus is considered ajunior synonym of Hemirrhagus
because they share characters of generic significance. Thus,
the subfamily Spelopelminae Smith, 1995 is here regarded as
asynonym of the subfamily Theraphosinae Thorell, 1870.
Also Cyrtopholis pernix (Ausserer, 1875) is transferred to
Hemirrhagus.Atotal of 15 species included in this genus are
diagnosed and keyed. Aphylogenetic analysis of the species
was performed and one fittest tree (fit=164.4; 47 steps; CI
0.55; RI 0.70) was found.
Introduction
The genus Hemirrhagus Simon, 1903 was known from
the female holotype of the type species, H. cervinus
(Simon, 1891), and from H. peruvianus Chamberlin,
1916 and H. major Chamberlin, 1916. The last two
species are here removed from Hemirrhagus because
they lack the diagnostic characters of the genus. Pérez-
Miles (1998) transferred Hemirrhagus from Ischnocoli-
nae incertae sedis,where it was placed by Raven (1985),
to Theraphosinae, on the basis of the presence of anew
type of urticating hairs found in the holotype of H.
cervinus.The study of types and additional material
attributed to Spelopelma Gertsch, 1982 led us to observe
that several species of this genus, as well as Cyrtopholis
pernix (Ausserer, 1875) and other undescribed species of
Hemirrhagus also have this type VI of urticating hairs.
Also the palpal organ morphology, the spermathecal
structure and the presence of retrolateral-ventral coxal
heels are shared by all the above-mentioned taxa, which
lead us to establish the synonymy of Spelopelma and
C. pernix with Hemirrhagus. Spelopelma was considered
as Theraphosidae incertae sedis by Raven (1985: 116).
Smith (1995) proposed the new subfamily Spelopelminae
for Spelopelma because of the weakly-developed eye
tubercle and eyes which may be completely absent in
some cave-dwelling species. Schmidt (1997) agreed with
this author but misinterpreted the urticating hairs as
being type II, as had also been done by Gertsch (1982:
91) for Spelopelma elliotti (Gertsch, 1973). Schmidt
(1997) also disregarded the presence of aclear keel on
the male palpal organ to support this conclusion. As a
consequence of our study the genus Hemirrhagus (in-
cluding its junior subjective synonym Spelopelma)ishere
placed in the Theraphosinae, mainly because of the
presence of type VI urticating hairs (of Pérez-Miles,
1998) and the presence of apalpal organ with an
extended subtegulum and adeveloped keel. The absence
of urticating hairs in some species could be interpreted
as reversions related to troglobitic habits, like the loss of
eyes. Thus the subfamily Spelopelminae, based solely on
the reduction or lack of eyes, cannot be supported, and
is here considered as ajunior synonym of Thera-
phosinae. Six new species of Hemirrhagus are here
described from Mexico. Aphylogenetic analysis of the
species resulted in one fittest tree of 47 steps, CI 0.55, RI
0.70, fit 164.4.
Methods
Abbreviations: AME=anterior median eyes, ALE=
anterior lateral eyes, PME=posterior median eyes,
PLE=posterior lateral eyes, OQ=ocular quandrangle
(including lateral eyes); d=dorsal, p=prolateral,
r=retrolateral, v=ventral; AMNH=American Museum
of Natural History, New York; BMNH=Natural
History Museum, London; LAAH=Laboratorio de
Acarología ‘‘Anita Ho mann’’, Facultad de Ciencias,
Universidad Autónoma de Mexico; LAE=Laboratorio
de Acarología, Instituto de Ecología, Universidad
Autónoma de Mexico; MCZ=Museum of Comparative
Zoology, Harvard, USA; MNHNP=Museum National
d’Histoire Naturelle, Paris, France; UNAM=
Universidad Autónoma de Mexico; USP=Universidad
de São Paulo, Brazil. All measurements are in mm.
Taxa:Fifteen species of Hemirrhagus considered in
the present review are included as terminal taxa for the
analysis. In afirst run the genera Euathlus Ausserer,
Grammostola Simon, Plesiopelma Pocock and Homo-
eomma Ausserer were used as outgroups to test the
monophyly of Hemirrhagus,considering their basal pos-
ition in the cladograms proposed by Pérez-Miles et al.
(1996) and Pérez-Miles (2000) for Theraphosinae.
Characters:Character distribution is shown in Table
10. ( 0 )Eyes: eight eyes present=0,PME absent=1,all
eyes absent=2.(1 )Ocular tubercle: normally developed
(as in general in theraphosids)=0,reduced=1,very
reduced (without elevation)=2.(2 )Periocular pigmen-
tation: absent=0,divided (involving some areas around
eyes)=1,entire (one area involving all eyes)=2.(3 )
Labial cuspules: numerous (more than 25)=0,reduced
(5–15)=1,very reduced (1 or 2)=2.(4 )Scopulae on tarsi
IV: divided by longitudinal band of longer and thicker
spiniform setae=0,entire=1.(5 )Female tibia I: with 0–8
spines=0,with 10 or more spines=1.(6 )Female tibia II
spination (coded as 5). ( 7 )Female tibia III spination
(coded as 5). ( 8 )Female tibia IV spination (coded
as 5). ( 9 )Female metatarsus Ispination (coded as 5).
( 10)Female metatarsus II spination (coded as 5).
365
( 11)Female metatarsus III spination (coded as 5). ( 12)
Female metatarsus IV spination (coded as 5). ( 13)Type
VI urticating hairs: absent=0,present=1.(14)Patch of
urticating hairs: di use, extended or oval=0,inaheart-
like patch=1,inabutterfly-like patch=2 (absence or
two patches coded as ?). ( 15)Disposition of urticating
hairs: one patch or di use=0,two round patches clearly
delimited=1 (absence coded as ?). ( 16)Area of urticating
hairs: not extended=0,extended laterally, in anterior–
posterior direction=1.(17)Spermathecal receptacles:
separated or slightly fused (less than half spermathecal
length)=0,widely fused (fusion zone more than half
spermathecal length)=1.(18)Spermathecal receptacles:
not close together at base (most species)=0,close to-
gether at base ( H. puebla)=1.(19)Spermathecal recep-
tacles: without clear inflexion in neck=0,with clear
inflexion in neck=1.(20)Subtegulum: not notched at
base of keel=0,notched at base of keel=1.(21)Special-
ised plumose setae on prolateral face of trochanter I:
absent=0,present=1.(22)Retrolateral–ventral coxal
heels: absent=0,present=1.Multistate character 2was
coded as non-additive.
Computer assisted methods:Adata matrix of 19 taxa
(including the outgroups) and 23 characters (Table 10)
was analysed. To test the monophyly of Hemirrhagus a
first run was carried out using an Ms-Dos computer
program, Hennig 86, developed by Farris (1988),
considering Euathlus, Grammostola, Plesiopelma and
Homoeomma as outgroups. Then, acladistic analysis
was performed using an Ms-Dos program, Pee-Wee,
developed by Golobo (1993a, b). To find the fittest
trees we used heuristic search with 15 additive sequences
and tree bisection reconnection; the concavity index was
6, following Ramírez (1998).
Genus Hemirrhagus Simon, 1903
Hemirrhagus Simon, 1903: 926; Strand, 1907: 16; 1912: 175;
Petrunkevitch, 1911: 71; 1928: 78; Roewer, 1942: 231; Raven,
1985: 116; Smith, 1995: 185; Pérez-Miles, 1998: 121.
Spelopelma Gertsch, 1982: 87; Smith, 1995: 32. New synonymy.
Type species: Hemirrhagus cervinus (Simon, 1891).
Diagnosis:Di ers from other theraphosine genera
by the retrolateral projections ventrally on coxae of all
legs. Also di ers by the presence of type VI urticating
hairs (Pérez-Miles, 1998) in most species, with the excep-
tion of H. grieta, H. mitchelli, H. puebla, H. reddelli and
H. stygius.Additionally di ers from most theraphosine
genera in the palpal organ morphology, with only one
retrolateral spiral keel (Figs. 5, 6, 14, 15, 24, 25). The
only other theraphosine genus which includes some
species having only one retrolateral keel on the palpal
organ is Aphonopelma Pocock, 1901, but in this genus
the keel is less developed and placed more distally and
prolaterally.
Composition:Fifteen species: Hemirrhagus cervinus
(Simon, 1891), H. chilango sp. n., H. coztic sp. n., H.
eros sp. n., H. gertschi sp. n., H. ocellatus sp. n.,
H. papalotl sp. n., H. elliotti (Gertsch, 1973) comb. n.,
H. grieta (Gertsch, 1982) comb. n., H. mitchelli
(Gertsch, 1982) comb. n., H. nahuanus (Gertsch, 1982)
comb. n., H. puebla (Gertsch, 1982) comb. n., H. reddelli
(Gertsch, 1973) comb. n., H. stygius (Gertsch, 1971)
comb. n., H. pernix (Ausserer, 1875) comb. n.
Misplaced species:Examination of type specimens of
Hemirrhagus major Chamberlin, 1916 and H. peruvianus
Chamberlin, 1916 (all deposited in MCZ, examined)
confirms that neither of these species belongs to
Hemirrhagus. Hemirrhagus major is formally transferred
to Hapalotremus Simon, 1903 comb. n., and H. peruvi-
anus is transferred to Homoeomma Ausserer, 1871
comb. n.
Comments: Spelopelma was placed in the Thera-
phosidae incertae sedis by Raven (1985); Smith (1995)
created the subfamily Spelopelminae for this genus,
based only on the reduction or absence of eyes, which
is usually found in troglobitic fauna. This was followed
by Schmidt (1997), who misinterpreted the urticating
hairs as being type II, as also did Gertsch (1982), and
disregarded the presence of adeveloped keel on the
palpal bulb. Spelopelma and Hemirrhagus share the
retrolateral–ventral heel on all coxae, the similar
morphology of the palpal organ, and the presence
of type VI urticating hairs in some species of both
genera. The absence of these urticating hairs in some
species seems to be asecondary loss related to the
adaptation to troglobitic habits, as is the loss or reduc-
tion of the eyes and ocular tubercle. Thus, the synonymy
between Spelopelma Gertsch, 1982 and Hemirrhagus
Simon, 1891, and that between Spelopelminae Smith,
1995 and Theraphosinae Thorell, 1870, is here estab-
lished. Cyrtopholis pernix (Ausserer, 1875) is here
transferred to Hemirrhagus because of sharing
the presence of type VI urticating hairs, palpal
organ morphology and retrolateral–ventral coxal
heels. Hemirrhagus major is placed in Hapalotremus
because of the characteristic palpal organ morphology
and other characters of generic significance. Hemirrha-
gus peruvianus is placed in Homoeomma because
of sharing the palpal organ morphology, with a
digitiform apophysis, and other characters of generic
significance.
Key to species
1. Urticating hairs present .......................................................... 2
—Urticating hairs absent ........................................................... 11
2. Urticating hairs arranged in two lateral round patches ......... 3
—Urticating hairs arranged otherwise ....................................... 4
3. Reduced number of labial cuspules (fewer than 15) .....................
.................................................................................. H. ocellatus
—Numerous labial cuspules (more than 25) ............... H. chilango
4. Periocular pigmentation divided (Fig. 22) in combination
with spinose metatarsi III and IV (more than 15 spines)
and divided scopula. Males with spinose tibia IV (10 spines
or more) .................................................................... H. gertschi
—Not as above .......................................................................... 5
5. Numerous labial cuspules (more than 25) ............................... 6
—Reduced number of labial cuspules (fewer than 15) .............. 9
6. Reduced spination on tibia IV (fewer than 5spines),
scopula entire, heart-like urticating hair patch .............. H. eros
—Normal spination on tibia IV (more than 15 spines) ............ 7
366
Revision of Hemirrhagus
7. Ocular tubercle reduced ......................................... H. nahuanus
—Ocular tubercle normally developed ...................................... 8
8. Plumose setae present on trochanter I(retrolateral face) .............
...................................................................................... H. pernix
—Plumose setae absent ................................................ H. cervinus
9. Tarsal scopula entire, butterfly-like urticating hair patch .............
................................................................................... H. papalotl
—Tarsal scopula divided, urticating hair patch otherwise ........ 10
10. Periocular pigmentation entire (surrounding all eyes, Fig. 8).......
....................................................................................... H. coztic
—Periocular pigmented area divided ............................. H. elliotti
11. Eight eyes present ................................................................... 12
—PME absent or all eyes absent ............................................... 14
12. Ocular tubercle well developed .................................. H. stygius
—Ocular tubercle reduced ......................................................... 13
13. Spermathecal receptacles widely fused, with narrow base
(Gertsch, 1982; fig. 16) ............................................... H. puebla
—Spermathecal receptacles separated ........................... H. reddelli
14. Eyes completely absent ................................................ H. grieta
—Only PME absent .................................................... H. mitchelli
Hemirrhagus cervinus (Simon, 1891) (Table 1)
Cratorrhagus cervinus Simon, 1891: 330 (D\ ); F. O. P.-Cambridge,
1899: 41, pl. 2, fig. 1.
Hemirrhagus cervinus:Simon, 1903: 926, fig. 1077; Strand, 1907: 16;
1912: 175; Petrunkevitch, 1911: 71; 1928: 78; Roewer, 1942:
231; Raven, 1985: 116; Smith, 1995: 185, figs. 1032–1033;
Pérez-Miles, 1998: 121, figs. 1–6.
Type:Holotype \ from Mexico, without locality data,
deposited in MNHNP, #756, examined.
Diagnosis:Di ers from most species of Hemirrhagus
by the presence of anormally developed ocular tubercle,
from H. pernix by the absence of plumose setae on
trochanter I, from H. eros by the divided tarsal scopulae
and from H. stygius by the presence of urticating hairs.
Female:Total length, not including chelicerae or
spinnerets, 32. Prosoma length 10.5, width 10. Anterior
eye row straight, posterior row recurved. Eye sizes and
interdistances: AME 0.76, ALE 0.40, PME 0.40, PLE
0.24; AME-AME 0.24, AME-ALE 0.10, PME-PME
0.76, PME-PLE 0.06, ALE-PLE 0.12; OQ length 1.00,
width 1.95, clypeus height 0.30. Ocular tubercle nor-
mally developed, periocular pigmentation absent. Fovea
transverse, straight, width 2.2. Labium length 1.2, width
1.7, with 35 cuspules. Maxillae with 126 cuspules.
Sternum length 4.7, posterior sternal sigilla large, oval,
submarginal. Chelicera with 20 teeth on promargin (8
small, 2medium, 10 large). Tarsi I–IV scopulate,
scopula divided. Metatarsi I–III scopulate, IV scopulate
on distal end. Length of legs and palpal segments as in
Table 1. Spination: femora II 1d; III 3d; IV 1d, 1p;
patellae I–IV 0; tibiae I6v, 2p; II 6v, 1p; III 7v, 3p, 3r;
IV 2v, 7p, 5r; metatarsi I3v; II 5v, 2p; III 3d, 6v, 4p, 4r;
IV 5d, 3v, 7p, 4r; tarsi I–IV 0. Prosoma and legs
orange-brown, opisthosoma yellow-brown with round
posterior grey-brown patch. Type VI urticating hairs
present in one patch. Spermathecae, see Pérez-Miles
(1998: fig. 6).
Distribution:Mexico, known only from the holo-
type.
Hemirrhagus chilango,new species (Figs. 1–6, Table 2)
Type:Holotype _ from Pedregal de San Ángel,
Mexico, D.F. (19( 18# N, 99( 11# W), 15 November 1977,
A. Zaldivar, deposited in LAAH.
Etymology:Noun in apposition, masculine singular.
Chilango is apopular name given to the people living in
Mexico City; it refers to the type locality.
Diagnosis:Di ers from other Hemirrhagus species
by the two oval patches of urticating hairs, situated
laterally and widely separated on the opisthosoma, and
from H. ocellatus by the more developed ocular tubercle
and larger number of labial cuspules (more than 25).
Male (Figs. 1–6): Total length, not including cheli-
cerae or spinnerets, 21.3. Prosoma length 9.1, width 8.3.
IIIIII IV
Femur 9.2 8.2 8.0 10.5
Patella 5.6 5.1 4.5 4.7
Tibia 7.0 6.3 6.0 8.3
Metatarsus 6.4 6.2 7.7 11.0
Tarsus 5.2 5.2 5.4 6.4
Table 1: Hemirrhagus cervinus (Simon). Female holotype, length of
leg segments (the holotype lacks palps).
Figs. 1–6: Hemirrhagus chilango sp. n., male holotype. 1 Body, dorsal
view; 2 Ocular region, dorsal view; 3 Prosoma, ventral
view;
4 Left palpal organ, ventral view; 5 Ditto, retrolateral
view;
6 Tibial apophysis of left leg I, prolateral view. Scale
lines=10mm(1, 3), 5mm(6), 1mm(2, 4, 5).
367
F. Pérez-Miles &A.Locht
Anterior eye row slightly procurved, posterior row re-
curved. Eye sizes and interdistances: AME 0.20, ALE
0.28, PME 0.14, PLE 0.30; AME-AME 0.20, AME-
ALE 0.16, PME-PME 0.76, PME-PLE 0.08, ALE-PLE
0.20; OQ length 0.64, width 1.54; clypeus height 0.16.
Ocular tubercle slightly reduced, periocular pigmen-
tation entire (Fig. 2). Fovea transverse, procurved,
width 1.60. Labium length 1.10, width 1.30, with 28
cuspules. Maxilla with 114 cuspules. Sternum length 3.6,
posterior sternal sigilla oval, submarginal (Fig. 3). Che-
licera with 15 teeth on promargin (5 small, 10 large).
Tarsi I–IV scopulate, scopula Ientire, II–IV divided.
Metatarsi Iscopulate, II–IV scopulate on distal end.
Tibia of leg Iwith prolateral double apophysis (Fig. 6).
Palpal organ with one wide keel (Figs. 4–5). Length of
legs and palpal segments as in Table 2. Spination:
femora II 4p; III 1d, 2r; patellae II 3p; IV 1d; tibiae I6v,
3p, 5r; II 7v, 5p, 4r; III 4v, 4p, 1r; IV 3v, 9p; palp 3v, 1p;
metatarsi I3v, 1p; II 2d, 3v, 2p, 1r; III 5v, 2p, 4r; IV 3d,
6v, 7p, 5r; tarsi I–IV 0. Prosoma and legs orange-brown,
opisthosoma yellow-brown with two golden lateral
patches. Type VI urticating hairs present, arranged in
two lateral patches (Fig. 1).
Female:Unknown.
Other material examined:Pedregal de San Ángel,
Mexico, D.F., 13 September 1985, 1juv., no collector’s
name, deposited in LAE.
Distribution:Known only from Pedregal de San
Ángel, Mexico, D.F.
Hemirrhagus coztic,new species (Figs. 7–10, Table 3)
Type:Holotype \ from Cueva del Diablo (19( 01# N,
99( 04# W), Tepoztlán, Morelos, Mexico, 26 November
1978, E. López, deposited in LAAH.
Etymology:Coztic is anoun in apposition taken
from the Nahuatl language and means yellow, referring
to the colour of this species.
Diagnosis:Di ers from most Hemirrhagus species
by the periocular pigmented area involving all eyes, from
H. ocellatus and H. chilango by the presence of one area
of urticating hairs extended laterally, and from H. pernix
by the smaller number of labial cuspules.
Female (Figs. 7–10): Total length, not including
chelicerae or spinnerets, 20.4. Prosoma length 9.3, width
7.6. Anterior eye row procurved, posterior row recurved.
Eye sizes and interdistances: AME 0.20, ALE 0.36, PME
0.04, PLE 0.30; AME-AME 0.16, AME-ALE 0.02,
PME-PME 0.58, PME-PLE 0.04, ALE-PLE 0.08; OQ
length 0.60, width 1.32; clypeus height 0.22. Ocular
tubercle slightly reduced, periocular pigmentation
entire (Fig. 8). Fovea transverse, procurved, width 1.00.
Labium length 1.5, width 1.1, with 12 cuspules. Maxilla
with 135 cuspules. Sternum length 4.0, posterior sternal
sigilla circular, submarginal (Fig. 10). Chelicera with 12
teeth on promargin (4 small, 8large). Tarsi I–IV scopu-
late, scopula I–II entire, III–IV divided. Metatarsi I–II
scopulate, III–IV scopulate on distal half. Length of legs
and palpal segments as in Table 3. Spination: femora I
1d; II 3d; III 3d; IV 2d; patellae I–IV 0; tibiae I6v, 1p;
II 6v, 2p; III 7v, 2p, 3r; IV 3v, 4p, 4r; metatarsi I2v; II
2v, 1p; III 8v, 3p, 3r; IV 8v, 10p, 3r; tarsi I–IV 0.
Prosoma and legs yellow-brown, opisthosoma grey-
brown with two golden lateral bands. Type VI urticating
hairs present, arranged in one patch extended laterally
(Fig. 7). Spermathecae as in Fig. 9.
Male:Unknown.
Other material examined:Cueva de San Juan,
Tepoztlán, Morelos, Mexico (19( 00# 12$ N, 99( 05# 12$ W),
16 November 1978, 1 \ 1juv., R. Castellanos, deposited
in LAE.
IIIIII IV Palp
Femur 8.3 7.9 7.5 9.5 6.1
Patella 4.6 4.2 3.8 4.1 3.5
Tibia 6.7 6.2 6.3 7.8 5.1
Metatarsus 7.3 6.9 7.1 11.5
Tarsus 5.9 5.4 5.5 6.6 2.5
Table 2: Hemirrhagus chilango sp. n. Male holotype, length of leg
and palpal segments.
Figs. 7–10: Hemirrhagus coztic sp. n., female holotype. 7 Body, dorsal
view; 8 Ocular region, dorsal view; 9 Spermathecae,
dorsal view; 10 Prosoma, ventral view. Scale
lines=10mm(7), 5mm(10), 1mm(8, 9).
IIIIII IV Palp
Femur 8.7 8.4 6.9 9.0 6.1
Patella 4.6 4.4 4.0 4.5 3.5
Tibia 7.8 7.0 6.8 8.7 5.0
Metatarsus 7.0 6.5 7.5 11.4
Tarsus 5.3 5.5 5.3 6.0 5.0
Table 3: Hemirrhagus coztic sp. n. Female holotype, length of leg and
palpal segments.
368
Revision of Hemirrhagus
Distribution:Known only from Tepoztlán, Morelos,
Mexico.
Hemirrhagus eros,new species (Figs. 11–20, Tables 4–5)
Types:Holotype _ and paratype \ from El Punto,
Oaxaca, Mexico (16( 19# N, 96( 28# W), 28 June 1967 and
5September 1962 respectively, both collected by M. R.
Bogert, deposited in AMNH.
Etymology:Named after the Greek god Eros, refer-
ring to the heart-like patch of urticating hairs with the
urticating hairs representing arrows, symbols of that
god.
Diagnosis:Di ers from most Hemirrhagus species
by the entire tarsal scopulae, and from H. papalotl by the
urticating hairs arranged in aheart-shaped patch. Also
di ers from other species by the very sparse spination on
tibiae and metatarsi III–IV.
Male (Figs. 11–16): Total length, not including che-
licerae or spinnerets, 26.0. Prosoma length 13.7, width
12.5. Anterior eye row slightly recurved, posterior row
recurved. Eye sizes and interdistances: AME 0.28, ALE
0.36, PME 0.26, PLE 0.38; AME-AME 0.30, AME-
ALE 0.14, PME-PME 1.14, PME-PLE 0.06, ALE-PLE
0.14; OQ length 0.84, width 1.56; clypeus height 0.34.
Ocular tubercle normally developed, periocular pigmen-
tation entire (Fig. 12). Fovea transverse, procurved,
width 1.40. Labium length 1.60, width 2.0, with 12
cuspules. Maxilla with 200 cuspules. Sternum length 5.4,
posterior sternal sigilla large, oval, submarginal (Fig.
13). Chelicera with 15 teeth on promargin (5 small, 10
large). Tarsi I–IV scopulate, scopula entire. Metatarsi
I–II scopulate, III scopulate on distal half, IV not
scopulate. Tibia of leg Iwith prolateral double apophy-
sis (Fig. 16). Palpal organ as in Figs. 14–15. Length of
legs and palpal segments as in Table 4. Spination:
femora I–IV and palp 0; patellae I–IV and palp 0; tibiae
I2v, 1p, 1r; II 1d, 3v, 2p, 2r; III 1p, 1r, 1v; IV 1v, 1p;
metatarsi II 3v; III 5v, 1p, 1r; IV 4v; tarsi I–IV 0.
Prosoma and legs dark brown, with light brown hairs on
carapace margin, opisthosoma dark brown with few
dark reddish hairs and platinum heart-shaped patch of
type VI urticating hairs in centre (Fig. 11).
Female (Figs. 17–20): Total length, not including
chelicerae or spinnerets, 35.0. Prosoma length 15.8,
width 14.5. Anterior eye row slightly procurved, pos-
terior row recurved. Eye sizes and interdistances: AME
0.34, ALE 0.50, PME 0.24, PLE 0.4; AME-AME 0.32,
AME-ALE 0.10, PME-PME 0.90, PME-PLE 0.04,
ALE-PLE 0.16; OQ length 0.92, width 1.9; clypeus
height 0.40. Ocular tubercle normally developed, peri-
ocular pigmentation entire (Fig. 18). Fovea transverse,
Figs. 11–16: Hemirrhagus eros sp. n., male holotype. 11 Body, dorsal
view;
12 Ocular region, dorsal view; 13 Prosoma, ven-
tral view; 14 Right palpal organ, retrolateral view; 15
Ditto, prolateral view; 16 Tibial apophysis of left leg I,
prolateral view. Scale lines=10mm(11, 13), 5mm(16),
1mm(12, 14, 15).
Figs. 17–20: Hemirrhagus eros sp. n., female paratype. 17 Body,
dorsal view;
18 Ocular region, dorsal view; 19 Prosoma,
ventral view;
20 Spermathecae, dorsal view. Scale
lines=10mm(17, 19), 1mm(18, 20).
IIIIII IV Palp
Femur 11.6 11.4 10.0 11.8 8.0
Patella 6.6 6.1 6.0 5.6 4.9
Tibia 10.5 9.1 8.3 11.2 6.8
Metatarsus 9.4 9.0 10.1 14.0
Tarsus 7.4 7.3 7.3 8.2 3.5
Table 4: Hemirrhagus eros sp. n. Male holotype, length of leg and
palpal segments.
369
F. Pérez-Miles &A.Locht
recurved, width 2.0. Labium length 1.8, width 2.3, with
42 cuspules. Maxilla with 145 cuspules. Sternum length
6.5, posterior sternal sigilla large, oval, submarginal
(Fig. 19). Chelicera with 19 teeth on promargin (8 small,
11 large). Tarsi I–IV densely scopulate, scopula entire.
Metatarsi I–II scopulate, III scopulate on distal end, IV
not scopulate. Length of legs and palpal segments as in
Table 5. Spination: femur I1d; III 1d; patellae I–IV 0;
tibiae II 2v, 1p; III 2v; IV 1v; metatarsi I3v; II 3v; III 1v,
1p; IV 2d, 2p, 1r; tarsi I–IV and palp 0. Prosoma
cinnamon brown with light brown hairs on carapace
margin, legs and opisthosoma dark brown with few dark
reddish hairs and platinum heart-shaped patch of type
VI urticating hairs (Fig. 17). Spermathecae as in Fig. 20.
Other material examined:ElPunto, Oaxaca,
Mexico, 5September 1962, 1adult \ ,28June 1967, 2
small \ ,all collected by C. M. Bogert, deposited in
AMNH.
Distribution:Known only from El Punto, Oaxaca,
Mexico.
Hemirrhagus gertschi,new species (Figs. 21–30, Tables
6–7)
Types:Holotype _ and paratype \ from Resumi-
dero, Guerrero, Mexico (18( 35# 52$ N, 99( 42# 40$ W), 27
March 1976, L. Rendon, deposited in LAAH.
Etymology:Named after the late American arach-
nologist, W. J. Gertsch, who contributed much to thera-
phosid taxonomy, particularly to the discovery of cave
theraphosids.
Diagnosis:Males di er from the other species by the
presence of numerous (11) spines on the dorsal surface
of tibia IV. Females di er from most species by the
periocular pigmentation involving only the anterior
eyes, from H. papalotl by the divided tarsal scopulae,
and from H. elliotti by the spinose metatarsi III and IV.
Male (Figs. 21–26): Total length, not including che-
licerae or spinnerets, 21.4. Prosoma length 9.6, width
8.8. Anterior eye row slightly recurved, posterior row
recurved. Eye sizes and interdistances: AME 0.22, ALE
0.20, PME 0.12, PLE 0.20; AME-AME 0.22, AME-
ALE 0.10, PME-PME 0.60, PME-PLE 0.04, ALE-PLE
0.14; OQ length 0.46, width 1.20; clypeus height 0.22.
Ocular tubercle very reduced, periocular pigmentation
surrounding AME only (Fig. 22). Fovea transverse,
recurved, width 1.0. Labium length 1.10, width 1.3, with
9cuspules. Maxilla with 64 cuspules. Sternum length
3.7, posterior sternal sigilla oval, submarginal (Fig. 23).
Chelicera with 18 teeth on promargin. Tarsi I–IV scopu-
late, scopula I–II entire, III–IV divided. Metatarsi I–III
scopulate, IV scopulate on distal half. Tibia of leg Iwith
prolateral double apophysis (Fig. 26). Palpal organ with
posterior retrolateral keel (Figs. 24–25). Length of legs
and palpal segments as in Table 6. Spination: femora I
2d; II 7d, 1p; III 4d; IV 2d, 2p; palp 4d; patellae I1v; II
1v; palp 1v; tibiae I4v, 2p, 4r; II 5v, 3p, 2r; III 5d, 6v, 3p,
5r; IV 11d, 6v, 3p, 5r; palp 6v, 3p; metatarsi I4v, 1p, 1r;
II 3v, 4p, 1r; III 3d, 3v, 3p, 2r; IV 1d, 5v, 6p, 4r; tarsi
I–IV 0. Prosoma and legs pale orange-brown, opistho-
soma yellow-brown with two golden lateral bands.
Opisthosoma and legs with yellow-brown setae. Type VI
urticating hairs present, arranged in two lateral bands
(Fig. 21).
IIIIII IV Palp
Femur 11.6 11.0 10.3 13.2 9.0
Patella 7.1 6.9 5.8 7.0 5.8
Tibia 8.2 7.2 7.1 9.2 6.0
Metatarsus 7.0 7.7 9.1 13.0
Tarsus 6.3 6.0 6.3 8.0 6.6
Table 5: Hemirrhagus eros sp. n. Female paratype, length of leg and
palpal segments.
Figs. 21–26: Hemirrhagus gertschi sp. n., male holotype. 21 Body,
dorsal view; 22 Ocular region, dorsal view; 23 Prosoma,
ventral view;
24 Left palpal organ, retrolateral view; 25
Ditto, prolateral view; 26 Tibial apophysis of left leg I,
prolateral view. Scale lines=5 mm (21), 1mm(22–26).
IIIIII IV Palp
Femur 11.1 11.0 10.0 12.2 7.4
Patella 5.5 5.3 4.5 5.2 4.4
Tibia 11.0 10.0 9.5 11.7 6.4
Metatarsus 10.6 10.3 11.5 15.6
Tarsus 8.2 7.6 7.6 8.9 3.3
Table 6: Hemirrhagus gertschi sp. n. Male holotype, length of leg and
palpal segments.
370
Revision of Hemirrhagus
Female (Figs. 27–30): Total length, not including
chelicerae or spinnerets, 27.0. Prosoma length 10.5,
width 9.2. Anterior eye row slightly recurved, posterior
row recurved. Eye sizes and interdistances: AME
0.22, ALE 0.30, PME 0.14, PLE 0.18; AME-AME 0.28,
AME-ALE 0.10, PME-PME 0.68, PME-PLE 0.06,
ALE-PLE 0.24; OQ length 0.50, width 1.40; clypeus
height 0.30. Ocular tubercle very reduced, periocular
pigmentation divided, surrounding anterior eyes (Fig.
28). Fovea transverse, recurved, width 1.21. Labium
length 1.20, width 1.6, with 15 cuspules. Maxilla with
121 cuspules. Sternum length 4.0, posterior sternal sigilla
large, oval, submarginal (Fig. 29). Chelicera with 20
teeth on promargin. Tarsi I–IV scopulate, scopula div-
ided. Metatarsi I–III scopulate, IV scopulate on distal
end. Length of legs and palpal segments as in Table 7.
Spination: femora I2d; II 2p; III 6d; IV 5d, 2p; palp 1p;
patellae I1v; palp 1v; tibiae I6v, 3p, 3r; II 9v, 4p, 5r; III
6v, 5p, 8r; IV 9v, 9r; palp 1v, 1r; metatarsi I2v; II 10v,
1p; III 2d, 8v, 5p, 6r; IV 10v, 5r; tarsi II 1v. Prosoma and
legs orange-brown, opisthosoma yellow-brown with two
gold lateral bands. Opisthosoma and legs with brown
setae. Type VI urticating hairs present, arranged in two
lateral bands (Fig. 27). Spermathecae as in Fig. 30.
Other material examined:Resumidero, Guerrero,
Mexico, without further data, 1 _ ,deposited in LAAH.
Distribution:Known only from the cave
Resumidero, Guerrero, Mexico.
Hemirrhagus ocellatus,new species (Figs. 31–34, Table 8)
Type:Holotype \ from Cueva Pen˜a Blanca, Valle
del Bravo, Estado de Mexico, Mexico (19( 11# N,
100( 08# W), 6November 1976, S. Ibarra, deposited in
LAAH.
Etymology:The species is named from the Latin
adjective ocellatus meaning ‘‘eyed’’, referring to the two
circular patches of urticating hairs on the abdomen.
Diagnosis:Di ers from most Hemirrhagus species
by the presence of two lateral circular patches of urti-
cating hairs, and from H. chilango by having the ocular
tubercle more reduced and fewer labial cuspules (fewer
than 15).
Female (Figs. 31–34): Total length, not including
chelicerae or spinnerets, 23.8. Prosoma length 10.0,
width 8.6. Anterior eye row procurved, posterior row
recurved. Eye sizes and interdistances: AME 0.20, ALE
0.32, PME 0.16, PLE 0.30; AME-AME 0.26, AME-
ALE 0.14, PME-PME 0.62, PME-PLE 0.08, ALE-PLE
0.14; OQ length 0.64, width 1.42; clypeus height 0.26.
Ocular tubercle very reduced, periocular pigmentation
entire (Fig. 32). Fovea transverse, recurved, width 1.58.
Figs. 27–30: Hemirrhagus gertschi sp. n., female paratype. 27 Body,
dorsal view;
28 Ocular region, dorsal view; 29 Prosoma,
ventral view;
30 Spermathecae, dorsal view. Scale
lines=5 mm (27), 1mm(28–30).
IIIIII IV Palp
Femur 10.5 10.6 9.2 11.4 7.1
Patella 5.9 5.0 4.3 4.8 4.5
Tibia 9.4 8.9 8.4 10.9 6.3
Metatarsus 8.2 8.5 10.4 14.3
Tarsus 6.4 6.4 6.1 7.7 6.0
Table 7: Hemirrhagus gertschi sp. n. Female paratype, length of leg
and palpal segments.
Figs. 31–34: Hemirrhagus ocellatus sp. n., female holotype. 31 Body,
dorsal view;
32 Ocular region, dorsal view; 33 Prosoma,
ventral view;
34 Spermathecae, dorsal view. Scale
lines=10mm(31), 5mm(33), 1mm(32, 34).
371
F. Pérez-Miles &A.Locht
Labium length 1.14, width 1.60, with 9cuspules. Maxilla
with 131 cuspules. Sternum length 4.4, posterior sternal
sigilla oval, submarginal (Fig. 33). Chelicera with 33
teeth on promargin (21 small, 12 large). Tarsi I–IV
scopulate, scopula I–II entire, III–IV divided. Metatarsi
I–II scopulate, III–IV not scopulate. Length of legs and
palpal segments as in Table 8. Spination: femora II 1d;
III 3d; IV 2d; patellae II 1v; tibiae I5v, 2p; II 9v, 4p; III
8v, 3p, 3r; IV 1d, 11v, 7p, 12r; metatarsi I5v, 1p; II 7v,
2p; III 2d, 7v, 5p, 4r; IV 2d, 10v, 6p, 3r; tarsi I–IV 0.
Prosoma and legs orange-brown, opisthosoma grey-
brown with two circular golden patches. Type IV urti-
cating hairs present, arranged in two circular lateral
patches (Fig. 31). Spermathecae as in Fig. 34.
Male:Unknown.
Distribution:Known only from Cueva Pen˜a Blanca,
Valle del Bravo, Estado de Mexico, Mexico.
Hemirrhagus papalotl,new species (Figs. 35–38, Table 9)
Type:Holotype \ from Gruta de Aguacachil
(18( 35# 30$ N, 99( 34# 29$ W), Taxco, Guerrero, Mexico,
29 November 1980, I. Cacaslera, deposited in LAAH.
Etymology:The specific epithet papalotl is taken
from the Nahuatl language, meaning butterfly, in
reference to the butterfly-like patch of urticating hairs.
Diagnosis:Di ers from most Hemirrhagus species
by the entire tarsal scopulae, and from H. eros by the
urticating hairs arranged in abutterfly-like patch, which
is placed more posteriorly than in H. eros.Also di ers
from H. eros by the presence of very numerous spines on
tibiae and metatarsi III–IV.
Female (Figs. 35–38): Total length, not including
chelicerae or spinnerets, 32.2. Prosoma length 12.7,
width 11.1. Anterior eye row straight, posterior row
recurved. Eye sizes and interdistances: AME 0.22, ALE
0.32, PME 0.18, PLE 0.26; AME-AME 0.26, AME-
ALE 0.16, PME-PME 0.78, PME-PLE 0.10, ALE-PLE
0.16; OQ length 0.64, width 0.64; clypeus height 0.40.
Ocular tubercle slightly reduced, periocular pigmen-
tation divided, involving only anterior eyes (Fig. 36).
Fovea transverse, recurved, width 0.25. Labium length
0.46, width 0.48, with 13 cuspules. Maxilla with 114
cuspules. Sternum length 5.8, posterior sternal sigilla
large, circular, submarginal (Fig. 37). Chelicera with 19
teeth on promargin (8 small, 11 large). Tarsi I–IV
scopulate, scopula entire. Metatarsi I–IV scopulate,
scopula divided. Length of legs and palpal segments as
in Table 9. Spination: femora I2d; II 4d; III 6d; IV 1d,
1p; patellae I2v; II 1v; tibiae I3d, 9v; II 3p, 7v; III 8v,
4p, 4r; IV 6v, 7p, 4r; metatarsi I4v; II 2v, 3p; III 1d, 8v,
3p, 4r; IV 1d, 12v, 5p, 4r; tarsi I–IV 0. Prosoma and legs
orange-brown, opisthosoma yellow-brown with pos-
terior golden patch of type VI urticating hairs in
butterfly-like form (Fig. 35). Spermathecae as in Fig. 38.
Male:Unknown.
Other material examined:Aguacachil, Guerrero,
Mexico: 3August 1980, 1 \ ,J.Palacios; 29 November
1980, 2juv., I. Cacaslera; 23 January 1982, 2juv., D.
Primero. All deposited in LAAH.
Distribution:All the material came from the same
cave (Gruta Aguacachil) in Taxco, Guerrero, Mexico.
Hemirrhagus elliotti (Gertsch, 1973), new combination
Schizopelma elliotti Gertsch, 1973: 144, fig. 2c (D\ ).
Spelopelma elliotti:Gertsch, 1982: 91, fig. 14; Schmidt, 1993: 68, fig.
130; Smith, 1995: 32, figs. 12–20.
Type:Holotype \ from Cueva de la Laguna
(24( 52# N, 100( 13# W), San Luis Potosi, Mexico,
deposited in AMNH, examined.
Diagnosis:Di ers from most Hemirrhagus species
by the combined presence of urticating hairs, reduced
ocular tubercle, and divided periocular pigmentation,
from H. gertschi by the reduced spination on female
IIIIII IV Palp
Femur 7.8 7.1 8.5 8.6 5.8
Patella 4.3 4.0 3.5 3.8 3.4
Tibia 6.6 5.6 6.0 7.7 4.7
Metatarsus 5.4 5.5 6.8 10.3
Tarsus 4.6 4.5 4.8 5.8 3.9
Table 8 Hemirrhagus ocellatus sp. n. Female holotype, length of leg
and palpal segments.
IIIIII IV Palp
Femur 12.6 11.8 11.5 13.6 8.5
Patella 6.5 6.5 5.4 5.5 5.2
Tibia 11.3 10.5 9.9 12.8 7.8
Metatarsus 11.2 11.0 10.8 17.0
Tarsus 8.7 8.0 7.6 8.9 7.1
Table 9: Hemirrhagus papalotl sp. n. Female holotype, length of leg
and palpal segments.
Figs. 35–38: Hemirrhagus papalotl sp. n., female holotype. 35 Body,
dorsal view;
36 Ocular region, dorsal view; 37 Prosoma,
ventral view;
38 Spermathecae, dorsal view. Scale
lines=10mm(35, 37), 1mm(36, 38).
372
Revision of Hemirrhagus
metatarsi III–IV (fewer than 10 spines), and from
H. papalotl by the divided tarsal scopulae.
Hemirrhagus grieta (Gertsch, 1982), new combination
Spelopelma grieta Gertsch, 1982: 93, fig. 17 (D\ ); Schmidt, 1993: 68,
fig. 133; Smith, 1995: 33, figs. 21–29.
Type:Holotype \ from Cueva de la Grieta
(17( 45# N, 97( 08# W), Huautla, Oaxaca, Mexico, de-
posited in AMNH, examined.
Diagnosis:Di ers from other Hemirrhagus species
by the complete absence of eyes (only subintegumental
spots present).
Hemirrhagus mitchelli (Gertsch, 1982), new combination
Spelopelma mitchelli Gertsch, 1982: 89, fig. 12 (D\ ); Schmidt, 1993: 68,
fig. 128; Smith, 1995: 34, figs. 30–40.
Type:Holotype \ from Entrada de Viento Alta
(22( 48# N, 98( 22# W), Tamaulipas, Mexico, deposited in
AMNH, examined.
Diagnosis:Di ers from other Hemirrhagus species
by the absence of PME (only subintegumental spots
present).
Hemirrhagus nahuanus (Gertsch, 1982), new combination
Spelopelma nahuanum Gertsch, 1982: 93, figs. 10–11 (D_ ); Schmidt,
1993: 68, figs. 126–127; Smith, 1995: 35, figs. 41–43.
Type:Holotype _ from Zoquitlán (18( 20# N,
97( 01# W), Puebla, Mexico, deposited in AMNH,
examined.
Diagnosis:Di ers from other Hemirrhagus species
by the very numerous labial cuspules, from H. cervinus,
H. pernix and H. eros by the reduced ocular tubercle,
and from H. chilango by the absence of two lateral oval
urticating hair patches.
Comments:Gertsch (1982) indicated the presence
of apatch of urticating hairs in this species, which
was also pointed out by Smith (1995). In our study we
did not find urticating hairs in the holotype, but
we presume they have been lost as aresult of
preservation conditions and manipulation in previous
studies. We therefore coded this character as present in
H. nahuanus.
Hemirrhagus puebla (Gertsch, 1982), new combination
Spelopelma puebla Gertsch, 1982: 91, fig. 16 (D\ ); Schmidt, 1993: 68,
fig. 132; Smith, 1995: 36, figs. 44–54.
Type:Holotype \ from Cueva de la Barranca,
Cuetzalan, Puebla, Mexico (20( 00# N, 97( 32# W),
deposited in AMNH, examined.
Diagnosis:Di ers from most Hemirrhagus species
by the presence of widely fused spermathecal
receptacles, and from H. elliotti by the absence of
urticating hairs.
Fig. 39: Cladogram of species of Hemirrhagus and outgroups (fit 164.4; 47 steps; CI 0.55; RI 0.70).
373
F. Pérez-Miles &A.Locht
Hemirrhagus reddelli (Gertsch, 1973), new combination
Schizopelma reddelli Gertsch, 1973: 143, fig. 2b (D\ ).
Spelopelma reddelli:Gertsch, 1982: 91, fig. 15; Schmidt, 1993: 68, fig.
131; Smith, 1995: 36, figs. 55–63.
Type:Holotype \ from Cueva del Nacimiento del
Rio San Antonio, Oaxaca, Mexico (16( 58# N, 95( 31# W),
deposited in AMNH, examined.
Diagnosis:Di ers from most Hemirrhagus species
by the absence of urticating hairs, and from H. puebla,
H. mitchelli, H. grieta and H. stygius by the periocular
pigmentation involving only PME.
Hemirrhagus stygius (Gertsch, 1971), new combination
Aphonopelma stygia Gertsch, 1971: 49 (D juv.).
Schizopelma stygia:Gertsch, 1973: 142, figs. 1, 2a (D\ ).
Schizopelma stygium:Brignoli, 1983: 140.
Spelopelma stygium:Gertsch, 1982: 89, fig. 13; Schmidt, 1993: 68, fig.
129; Smith, 1995: 37, figs. 64–65.
Type:Immature holotype from Cueva de los
Potrerillos, San Luis Potosi, Mexico (21( 19# N, 99( 03# W),
deposited in AMNH, examined.
Diagnosis:Di ers from most Hemirrhagus species
by the absence of urticating hairs, and from H. puebla,
H. mitchelli and H. grieta by the well-developed ocular
tubercle.
Hemirrhagus pernix (Ausserer, 1875), new combination
Crypsidromus pernix Ausserer, 1875: 178, pl. 6, figs. 22–23 (D_ ).
Hapalopus pernix:F.O.P.-Cambridge, 1897: 31, pl. 2, fig. 14.
Cyrtopholis pernix:Pocock, 1903: 98; Simon, 1903: 931; Smith, 1995:
182, figs. 1011–1021; Platnick, 1998: 158.
Type:Holotype _ from Pico de Orizaba, Veracruz,
Mexico (19( 01# N, 97( 16# W), deposited in BMNH,
examined.
Diagnosis:Di ers from other Hemirrhagus species
by the presence of specialised plumose setae on the
prolateral face of trochanter I.
Comments:This species is here removed from
Cyrtopholis because nodes on the male palpal tibiae and
type Iurticating hairs are absent. The palpal organ
morphology, coxal heels and the presence of type VI
urticating hairs led us to include this species in
Hemirrhagus.
Cladistic analysis
Using Hennig 86, two most parsimonious trees sup-
ported the monophyly of Hemirrhagus (188 steps, CI
0.85 and RI 0.90). Using Pee-Wee one fittest tree of 47
steps, CI 0.55, RI 0.70 and fit=164.4 was found (Fig.
39). Character fit and steps are given in Table 11. The
most obvious synapomorphy of the genus was the
retrolateral-ventral coxae with heels (character 22). An-
other probable additional synapomorphy of the genus is
the presence of only one wide keel on the palpal organ,
but this was not included in the analysis since only afew
males are known. The ingroup was dichotomously re-
solved on the basis of type VI urticating hairs (character
13); this seems to be asynapomorphy of apical
012345678910111213141516171819202122
Euathlus 00001????? ???0000001000
Grammostola 0000100000 001000000?000
Plesiopelma 00000001?0 0010000001000
Homoeomma 0000000000 0010000001000
H. cervinus 0000000110 011100?000?01
H. chilango 01200????? ???1?10???101
H. coztic 0121000110 0111001000?01
H. eros 0020100000 0001100000001
H. gertschi 0211011110 0111001000101
H. ocellatus 0221001110 ?111?10000?01
H. papalotl 0111111110 0111201000?01
H. elliotti 02110??11?? 001001100?01
H. grieta 22?20??11? ?110???000?01
H. mitchelli 12?00??11? ?110???000?01
H. nahuanus 02?00????? ???1000???001
H. puebla 02?100000?? ?10???111?01
H. reddelli 02?10?110?0 100???001?01
H. stygius 00?10????? ???0???000?01
H. pernix 00?00????? ?11100????111
Table 10: Data matrix used in the cladistic analysis.
Character 01 23456 78910111213141516171819202122
Fit 10 68.5 5.4 7.5 10 8.5 8.5 10 ——8.5 7.5 8.5 8.5 10 10 8.5 10 8.5 10 —10
Steps 263 73 12 21——2 323 112121—1
Extra steps 041 52 01 10——1 211 001010—0
Table 11: Character fit, steps and extra steps (the same values were found in the two trees).
374
Revision of Hemirrhagus
Hemirrhagus species with aparallelism in the basal
H. eros.Asfar as we know, this type of hair is unique in
Theraphosinae. The reduction of the ocular tubercle
(character 1), presumably related to troglobitic habits,
appears in the node of H. puebla with the apical clade,
with areversion in the clade of H. cervinus+ H. pernix.
We mapped the troglobitic habits on the cladogram and
involved most Hemirrhagus species, with ambiguities
in H. eros, H. nahuanus, H. chilango, H. cervinus and
H. pernix mainly because of missing entries in this
character.
Acknowledgements
We thank Dr Norman I. Platnick (AMNH) for the
loan of type specimens and for hospitality provided
while one of us (AL) worked in his laboratory, and
Laura Leibensperger (MCZ) for the loan of type speci-
mens. We also wish to thank Ignacio Vázquez, Dr Juan
Morales-Malacara and Dr Guadalupe Lopez Campos
(UNAM) for their help and hospitality during the study
visits of one of us (FPM). Adraft of the manuscript was
carefully read and criticised by Rogerio Bertani (USP).
We also thank two anonymous reviewers for their
helpful comments.
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F. Pérez-Miles &A.Locht
... Chamberlin (1916) described Hemirrhagus major Chamberlin, 1916 from the Cusco Valley, Peru based on a male and a female. Pérez-Miles & Locht (2003), whilst revising the genus Hemirrhagus Simon, 1903, found H. major to be misplaced, transferring it to Hapalotremus, based on male palpal bulb morphology. Over a decade later, Cavallo & Ferretti (2015) described a third species for the genus, Hapalotremus martinorum Cavallo & Ferretti, 2015 from both sexes. ...
... Hapalotremus Simon, 1903: 930. Hemirrhagus: Chamberlin (1916 Hapalotremus: Petrunkevitch (1911), Waterhouse (1912), Carbonell (1919), Carbonell (1923), Mello-Leitão (1923), Petrunkevitch (1928), Petrunkevitch (1939), Neave (1939), Roewer (1942), Bonnet (1957), Gerschman & Schiapelli (1973, Raven (1985), Schmidt (1986), Schmidt (1993aSchmidt ( , 1993b, Pérez-Miles et al. (1996), Schmidt (1997), Vol (1999), Schmidt (2002), Schmidt (2003), Pérez-Miles & Locht (2003), Cavallo & Ferretti (2015), Lucas & Indicatti (2015), Ferretti et al. (2018). ...
Article
Full-text available
The holotype of Hapalotremus albipes Simon, 1903 is redescribed. Four new species of the genus Hapalotremus Simon, 1903 are described: Hapalotremus apasanka sp. nov., Hapalotremus hananqheswa sp. nov., and Hapalotremus kaderkai sp. nov. from Peru, and Hapalotremus yuraqchanka sp. nov. from Bolivia. Discussion on palpal bulb morphology, ontogenetic spermathecal morphology, and biogeography is presented. Historical information about collectors and comments on the accession numbers and other label information of historical specimens is also presented.
... Mendoza (2014) in his taxonomic revision of Hemirrhagus described for the first time an adult male of one troglobitic tarantula: H. stygius. This species shares with other troglophilous and troglobitic Hemirrhagus the following troglomorphisms or evolutionary adaptations for life in caves: (a) the reduction of eyes, (b) legs long and slender, (c) lack of pigmentation and (d) urticantig setae type VI arranged in one dorsal or two paramedian patches (troglophiles) or absence of these in troglobites ( Pérez-Miles & Locht 2003;Mendoza, 2014). ...
... Type VI urticating setae were reported as exclusive of Hemirrhagus and can be arranged in one dorsomedian patch, two dorsal paramedian patches, or two lateral patches (Mendoza 2014). Only five species lack urticating setae associated with their high degree of troglomorphism ( Pérez-Miles 1998;Pérez-Miles & Locht 2003;García-Villafuerte & Locht 2010;Mendoza 2014;Estrada-Alvarez 2014). One of the most remarkable features found in this study was the presence of spinose setae dorsally on opisthosoma in the male of troglobitic H. akheronteus sp. ...
Article
Full-text available
The suborder Mygalomorphae is generally poorly represented in the cave faunas of the world. The genus Hemirrhagus is endemic to Mexico and has 22 described species. It is the only one with epigean, troglophile and troglobitic species. Lack of urticating setae, loss of eye pigmentation and ocular reduction are interpreted as evolutionary reversals related to their troglobitic habits. From five troglobitic species in the genus, only the male of Hemirrhagus stygius is known. Five new troglobitic species are described with both sexes: Hemirrhagus akheronteus sp. nov., Hemirrhagus billsteelei sp. nov., Hemirrhaugus diablo sp. nov., Hemirrhagus kalebi sp. nov. and Hemirrhagus sprousei sp. nov. The female of Hemirrhagus chilango is described for the first time. Hemirrhagus akheronteus sp. nov. has a group of spinose setae on the opisthosoma; this setal modification was never reported in any other theraphosid spiders. Females of Hemirrhagus kalebi sp. nov. and Hemirrhagus sprousei sp. nov. lay fixed hammock egg sacs, which is an unusual behavior among species in Theraphosinae. Two stridulating setae previously reported only in epigean species are present on some of the new troglobites.
... Several cave-dwelling species are bona fide troglobites that possess well-defined troglomorphies (e.g., reduction in size or complete loss of eyes, reduction of ocular pigmentation, elongation of appendages, loss of urticating setae). Future phylogenetic work (also see Pérez-Miles and Locht 2003) should help elucidate a variety of interesting biogeographic phenomena in this captivating genus. ...
Chapter
The Aviculariinae spiders sensu lato are known as the American arboreal tarantulas. They are characterized mainly by having legs with few or no spines, laterally extended tarsal and metatarsal scopulae, resulting in a spatulate appearance of the appendices, absence of spiniform setae on the prolateral maxillae, females with completely separated spermathecae, males with palpal bulb with subtegulum not extended, and long and thin embolus without keels (except Antillena). Some Aviculariinae, together with all Theraphosinae, are the only spiders that evolutionarily acquired urticating setae as a defense mechanism. The primary mechanism for releasing the urticating setae in Theraphosinae is by the friction of the legs with the abdomen, which throws the urticating setae into the air, in contrast, in most Aviculariinae the releasing mechanism occurs by direct contact. The Aviculariinae tarantulas have received considerable taxonomic and biological attention and the validity as a monophyletic group has been discussed extensively. Some phylogenetic studies suggest at least two subfamilies for the American arboreal tarantulas and their kin: Aviculariinae and Psalmopoeinae. Likewise, the phylogenetic relationships of these groups have been questioned, linking these tarantulas more closely with African or American taxa. Taxonomy, systematics and some aspects of its natural history, behavioral and distribution are addressed in this chapter.
... Several cave-dwelling species are bona fide troglobites that possess well-defined troglomorphies (e.g., reduction in size or complete loss of eyes, reduction of ocular pigmentation, elongation of appendages, loss of urticating setae). Future phylogenetic work (also see Pérez-Miles and Locht 2003) should help elucidate a variety of interesting biogeographic phenomena in this captivating genus. ...
Chapter
Studying morphology of Theraphosidae spiders can be very challenging, especially if the main objective is assembling characters for systematics. Such spiders present a homogeneous morphology, which, according to some specialists, has driven the attention of systematists to other groups of Araneae. Nevertheless, a great diversity of cuticular structures has been overlooked until the widespread use of scanning electron microscopy (SEM) in the last years for theraphosids. Among all mygalomorphs, Theraphosidae spiders possess the greatest variety of cuticular features. Data regarding cuticular features are still incipient, but we have been gathering massive quantity of SEM images of all parts of the spider body, revealing interesting structures to be used in systematics and investigated for functional morphology. In addition to the well-known tarsal adhesive setae of theraphosids and the urticating setae of Theraphosinae, we found putative chemosensitive setae, a great variety of stridulating setae, distinct morphologies of leg and palpal structures, including cuticular projections, labial and maxillary cuspules, trichobothria, as well as other enigmatic features. In this chapter, we aim to present a comprehensive revision of cuticular features of New World Theraphosidae spiders, with descriptions and micrographs.
... Together, the characters and character states make up character statements. Based on this definition, the coding of characters was carried out taking phylogenetic works carried out in other genera of the family as a starting point (Bertani, 2001;Fukushima, 2005;Pérez-Miles, 2000;Pérez-Miles & Locht, 2003;West et al., 2008;West & Nunn, 2010;Fukushima & Bertani, 2017). Intraspecific variation of each Brachypelma species was determined by reviewing between five and ten specimens of each sex for each species, with the exception of Central American species, because it was not possible to obtain additional samples to those already deposited in scientific collections. ...
Article
Full-text available
The tarantula genus Brachypelma includes colourful species that are highly sought after in the commercial pet trade. They are all included in CITES appendix II. We present phylogenetic analyses using molecular and morphological characters to revise Brachypelma, in which we include all currently known species. Our results agree with a previous study that shows the non-monophyly of Brachypelma. Both phylogenies strongly favour the division of Brachypelma into two smaller genera. The first clade (Brachypelma s.s.) is formed by B.albiceps, B. auratum, B. baumgarteni, B. boehmei, B. emilia, B. hamorii, B. klaasi and B. smithi. The species included in the second clade are transferred to the new genus Tliltocatl and is formed by T. albopilosum comb. nov., T. epicureanum comb. nov., T. kahlenbergi comb. nov., T. sabulosum comb. nov., T. schroederi comb. nov., T. vagans comb. nov. and T. verdezi comb. nov. Both genera can be differentiated by their coloration and the shape of the genitalia. We transfer to Tliltocatl: T. alvarezi, T. andrewi and T. aureoceps, but should be considered as nomina dubia. In addition, we transfer B. fossorium to Stichoplastoris. We discuss the implications of these taxonomical changes for CITES and for the Mexican Laws for wildlife protection.
... The aim of the revision is 3) also to critically evaluate what is known in this field of research and 4) to compare the evolutionary hypotheses of UrS development derived from the mentioned phylogenetic analyses. [14] Avicularia Lamarck, 1818 -+ ----- [15] Caribena Fukushima & Bertani, 2017 -+ ----- [14] Iridopelma Pocock, 1901 -+ ----- [15] Pachistopelma Pocock, 1901 -+ ----- [15] Typhochlaena C. L. Koch, 1850 -+ ----- [15] Ybyrapora Fukushima & Bertani, 2017 -+ ----- [14] PSALMOPOEINAE Ephebopus Simon, 1892 ----+ -- [5] Psalmopoeus Pocock, 1895 ------- [16] Pseudoclamoris Hüsser, 2018 ------- [17] Tapinauchenius Ausserer, 1871 ------- [6,16] THERAPHOSINAE Acanthoscurria Ausserer, 1871 + -+ ---- [18] Acentropelma Pocock, 1901 a + -? ---- [19,20] Aenigmarachne Schmidt, 2005 -----+ - [21] Agnostopelma Pérez-Miles & Weinmann, 2010 --+ + --- [22] Aguapanela Perafán & Cifuentes, 2015 --+ + --- [23] Aphonopelma Pocock, 1901 + ------ [6,18,24] Bistriopelma Kaderka, 2015 --+ ---- [25] Bonnetina Vol, 2000 --+ ---- [26] Brachypelma Simon, 1891 + -+ ---- [18] Bumba Chromatopelma Schmidt, 1995 --+ ---- [16] Citharacanthus Pocock, 1901 + ------ [18] Clavopelma Chamberlin, 1940 + ------ [16] Cotztetlana Mendoza Marroquín, 2012 + ------ [29] Crassicrus Reichling & West, 1996 + ------ [16] Cubanana Ortiz, 2008 + -+ ---- [30] Cyclosternum Ausserer, 1871 --+ ---- [18] Cyriocosmus Simon, 1903 --+ ---- [31,32] Cyrtopholis Simon, 1892 + -+ ----types I, III [6]; type III [16]; type I [18] Davus Cambridge, 1892 --+ ---- [16,19] Euathlus Ausserer, 1875 --+ + --- [33] Eupalaestrus Pocock, 1901 + -+ ---- [18] Eurypelmella Strand, 1907 a + -? ---- [19,20] Grammostola Simon, 1892 --+ + --- [6]; as Phrixotrichus in [18] Hapalopus Ausserer, 1875 b --+ + --- [18], [31] Hapalotremus Simon, 1903 --+ ---- [18] Hemirrhagus Simon, 1903 -----+ - [13,34,35] Homoeomma Ausserer, 1871 --+ + --- [18] Kankuamo Perafán et. al., 2016 ------+ [11] Kochiana Fukushima et al., 2008 --+ ---- [36] (Continued ) ...
Article
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Tarantula urticating setae are modified setae located on the abdomen or pedipalps, which represent an effective defensive mechanism against vertebrate or invertebrate predators and intruders. They are also useful taxonomic tools as morphological characters facilitating the classification of New World theraphosid spiders. In the present study, the morphology of urticating setae was studied on 144 taxa of New World theraphosids, including ontogenetic stages in chosen species, except for species with urticating setae of type VII. The typology of urticating setae was revised, and types I, III and IV were redescribed. The urticating setae in spiders with type I setae, which were originally among type III or were considered setae of intermediate morphology between types I and III, are newly considered to be ontogenetic derivatives of type I and are described as subtypes. Setae of intermediate morphology between that of body setae and type II urticating setae that were found in Iridopelma hirsutum Pocock, 1901 and Antillena rickwesti (Bertani & Huff, 2013) may provide another evidence that type II urticating setae evolved from body setae. It is supposed that the fusion of barbs with the shaft may lead to the morphology of type II setae. As the type II setae of Aviculariinae evolved independently to the UrS of Theraphosinae and both subfamilies represent two non-sister groups, this should explain the differences in the morphology of body setae in Aviculariinae and Theraphosinae. The terminology of “barbs” and “reversed barbs” was revised and redefined, newly emphasizing the real direction of barbs.
... Compared with Araneomorphae and other arachnid groups, few mygalomorph species with troglobite habits have been reported. They present the typical troglobite characteristics such as reduction or complete loss of eyes and pigmentation; some theraphosids from caves have also been shown to lose urticating setae (Pérez-Miles and Locht 2003). In the Neotropics, cave mygalomorphs such as Actinopus Karsch 1879, and Lasiodora C.L. Koch 1850 (Theraphosidae) have been described (Gertsch 1982;Goloboff 1994;Bernardi et al. 2007;de Armas et al. 2009;Bertani 2013;Mendoza 2014;Pedroso and Baptista 2014;Guadanucci et al. 2016;Guadanucci et al. in press). ...
Chapter
Full-text available
The infraorder Mygalomorphae comprises almost 3000 species. It is widely distributed all over the world, and about one third of the species included in this infraorder is represented in the Neotropics. The knowledge on several aspects of the biology of Mygalomorphae is relatively scarce in comparison with the Araneomorphae. However, recent studies developed all over the world, but especially in the Neotropical region, are continuously contributing to add information on foraging strategies, communication, reproductive biology, habitat selection, and defenses against natural enemies, minimizing the differences in the status of knowledge between these groups. In this chapter, we review some the main topics on Mygalomorphae biology, including behavior and ecology.
Chapter
Biogeography is a multidisciplinary field that is concerned with delimiting and explaining the geographic distributions of organisms in space and time. Due to their distribution patterns and interesting biology (e.g., ancient lineage with a nearly cosmopolitan distribution, sedentary lifestyle with limited dispersal capabilities), tarantulas are an appealing taxonomic group for addressing a variety of biogeographic questions concerning the Earth’s history. In this chapter, we discuss some biogeographic basic concepts, delve into the distribution patterns of New World tarantulas, and explore some of the historical explanations that may have led to these distributions. We mostly review and highlight the results of recent studies but also include personal observations and unpublished data. The distributions of higher-level taxonomic groups (subfamilies and tribes) are described and we detail their latitudinal and elevational limits. We also review the distributions of groups with unique insular habitats such as those found on islands surrounded by seas, forested “islands” surrounded by “seas” of deserts, and caves. Furthermore, we discuss the distribution of some unique morphological characters of taxonomic importance such as urticating setae. Finally, we review a handful of studies that have explicitly investigated the biogeography of New World tarantulas using a variety of different analytical methods.
Chapter
Tarantula urticating setae are modified setae located on the abdomen or pedipalps, which represent an effective defensive mechanism against vertebrate and invertebrate predators and intruders. They are also useful taxonomic tools as morphological characters facilitating the classification of New World theraphosid spiders. Up to now, seven morphological types of urticating setae, types I, II, III, IV, V, VI, and VII, have been described. They differ in location, shape, size, and orientation of barbs along the shaft and the length/width ratio. The urticating setae, which were originally among type III in spiders with type I setae, or were considered setae of intermediate morphology between types I and III, are now considered to be ontogenetic derivatives of type I. Recently published papers focused on the molecular phylogenetic reconstruction of Theraphosidae revealed the monophyly of theraphosine taxa with type I setae and the reclassification is also supported by the specific development of type I setae during ontogeny and by the morphological differences between type III setae in taxa with types I + III and types III or III + IV. The hypothesis of urticating setae evolution proposed and based on published molecular analyses gave proof that theraphosids with abdominal urticating setae of type I (Theraphosinae, part) and type II (Aviculariinae) had evolved independently and represent two non-sister but monophyletic groups. Although the taxa with urticating setae of types III and III + IV represent two monophyletic sister groups, the systematic position of Hemirrhagus with type VI setae and Kankuamo with type VII setae is still unclear, similarly to the evolution of both types of urticating setae.
Article
Full-text available
New specimens of the genus Hapalotremus (Theraphosinae) are revised based on the examination of types and additional material collected in Argentina, Bolivia and Peru. A new generic diagnosis is proposed. New information and illustrations are provided for known species and seven new species were recognized and are newly described and illustrated. Hence, Hapalotremus comprises 10 valid species, distributed along the Andes and Yungas in western South America. All species are keyed and mapped. New taxonomic features are included in the descriptions and the detail of embolus keels morphology and positions are described for the first time for the genus. Information on species habitat is included. The highest altitude record for a Theraphosidae spider, living at 4524 m above sea level, is reported. www.zoobank.org/urn:lsid:zoobank.org:pub:1C028676-CCD9-4654-A755-6ACA3BCE3E98
Article
Abstract— A new method for weighting characters according to their homoplasy is proposed; the method is non-iterative and does not require independent estimations of weights. It is based on searching trees with maximum total fit, with character fits defined as a concave function of homoplasy. Then, when comparing trees, differences in steps occurring in characters which show more homoplasy on the trees are less influential. The reliability of the characters is estimated, during the analysis, as a logical implication of the trees being compared. The “fittest” trees imply that the characters are maximally reliable and, given character conflict, have fewer steps for the characters which fit the tree better. If other trees save steps in some characters, it will be at the expense of gaining them in characters with less homoplasy.
L ocht Hemirrhagus reddelli (Gertsch, 1973), new combination Schizopelma reddelli Gertsch
  • F Pérez-Miles
F. Pérez-Miles &A.L ocht Hemirrhagus reddelli (Gertsch, 1973), new combination Schizopelma reddelli Gertsch, 1973: 143, fig. 2b (D\ ).
1875: 178, pl. 6, figs
  • Ausserer Crypsidromus Pernix
Crypsidromus pernix Ausserer, 1875: 178, pl. 6, figs. 22–23 (D_ ).
Zur Position der Gattung Spelopelma GertschAraneida: Theraphosidae)
SCHMIDT, G. E. W. 1997: Zur Position der Gattung Spelopelma Gertsch, 1982 (Araneida: Theraphosidae). Arachnol. Mag. 5 (8): 1–5.