Redefinition And Generic Revision Of The North American Vaejovid Scorpion Subfamily Syntropinae Kraepelin, 1905, With Descriptions Of Six New Genera
Author
González-Santillán, Edmundo
Author
Prendini, Lorenzo
text
Bulletin of the American Museum of Natural History
2013
2013-12-02
2013
382
1
71
http://www.bioone.org/doi/abs/10.1206/830.1
journal article
7646
10.1206/830.1
f86801d1-5089-42e6-9f91-423312e06eba
0003-0090
4611867
5793E88C-9654-41BC-BAF0-36017C1AEE95
Kochius
Soleglad and Fet, 2008
Figures 1B
,
2B
,
5
,
7
,
9–11
,
16C
,
18A, B
,
22A
,
23A
,
24A
,
25A
;
table 1
Buthus punctipalpi
Wood, 1983
[
=
Kochius punctipalpi
(Wood, 1983)
],
type
species, by original designation.
Vaejovis punctipalpi
group (part):
Williams, 1971a: 37
;
1971b: 44
;
1980: 80
;
Sissom and Francke, 1985: 1
;
Sissom, 1989: 180
;
1991: 26
;
Stockwell, 1992: 408
, 409;
Sissom, 1993: 68
;
Lourenço and Sissom, 2000: 135
;
Sissom, 2000: 532
, 548, 551;
Armas and Martín-Frías, 2001: 1
;
Hendrixson, 2001: 1
;
Ponce-Saavedra and Sissom, 2004: 539
, 541;
Sissom and Hendrixson, 2005a: 131
;
2005b: 35
;
Soleglad and Fet, 2005: 4
, 6, 7;
Graham and Fet, 2006: 7
, 8, 10;
Fet et al., 2006a: 9
,
table 1
; 2006b: 8; 2006c: 7;
Soleglad and Fet, 2006: 6
, 26,
tables 2
,
3
;
Graham and Soleglad, 2007: 11
, 12;
Francke and González-Santillán, 2007: 586
, 587, 590;
Soleglad et al., 2007: 134
, 135;
Soleglad and Fet, 2008: 1
, 26, 57, 60, 94, 95, 102;
McWest, 2009
: figs. 322–326,
tables 1
, 53, 89, 91, 93, 101– 105, 108, 123.
Kochius
Soleglad and Fet, 2008: 1
, 3, 34, 38, 51, 53, 54, 57, 58, 60–69, 71, 73, 74, 76, 79, 91–96, 102, 104, figs. 192, 193–196, 204,
tables 2
, 4, 6, 9 (part);
Ayrey and Soleglad, 2011: 1
(part).
DIAGNOSIS:
Kochius
differs from all other genera of
Syntropinae
in possessing the longest macrosetae on the telson vesicle of the female and, in some species, of both sexes (fig. 25A); some macrosetae are as long as or longer than the aculeus, becoming narrower and translucent terminally. Species of
Kochius
are also characterized by a raised, whitish boss, restricted to the posteromedial third of mesosomal sternite VII (fig. 23A). Although other genera of
Syntropinae
possess whitish glandular surfaces on sternite VII (fig. 23B), the structure in
Kochius
, which is noticeably more elevated and usually shaped like a truncate triangle, is unique. Most
Syntropinae
also display a glandular surface posteromedially on sternite V, but it is more pronounced in
Kochius
(fig. 23A). Additional diagnostic characters of
Kochius
are as follows. Species of
Kochius
possess more pronounced, granular carinae on the pedipalp chela manus, especially the drs, drsa, and rlm carinae (fig. 18A, B), and the pedipalp patellar rlds and rlm carinae are more coarsely granular than in other genera of
Syntropinae
. The pedipalp carinae of
Kochius
are also distinctive in comprising moniliform granules (fig. 18A, B), rather than clustered granules, as in other genera (fig. 17A–D). The patellar dorsal intercarinal surface (between the dpl and drl carinae) is flat in
Kochius
, compared to other genera in which it is slightly concave. The densely and coarsely granular (shagreened) carapace (fig. 16C), markedly shagreened mesosomal posttergites I–VII and sternite VII (fig. 23A), and uniformly finely granular (matte) ventral intercarinal surfaces of the pedipalp chela manus, retrolateral intercarinal surfaces of the pedipalp patella and intercarinal surfaces of metasomal segments I–V (fig. 24A) are also characteristic for the genus. Pedipalp chela trichobothrium
esb
is situated at RD6 and/or its associated macroseta, and trichobothrium
Db
is situated dorsal to the drl carina, compared with
Paravaejovis
, in which
esb
is situated between RD6 and RD5 (fig. 19D), and
Mesomexovis
,
gen. nov.
, in which
Db
is situated on the drl carina (fig. 19A). The macrosetae of legs I–IV also tend to be shorter and stouter in
Kochius
than in other genera (fig. 22A).
Four species of
Chihuahuanus
,
gen. nov.
(
C. cazieri
,
comb. nov.
,
C. crassimanus
,
comb. nov.
,
C. kovariki
,
comb. nov.
, and
C. russelli
,
comb. nov.
) resemble
Kochius
in possessing a raised, whitish boss in the posteromedial third of mesosomal sternite VII (fig. 23A), but may be separated from the latter by the small fusiform, whitish glandular area, anterior to the base of the aculeus, on the dorsal surface of the telson vesicle of adults; the shorter macrosetae on the telson vesicle of females; the less-developed (lower and smoother) pedipalp chelal rlm and rlvs carinae, and patellar rlds and rlm carinae; and the clustered instead of moniliform granules of the pedipalp carinae.
Species of
Thorellius
resemble
Kochius
with respect to the incrassate pedipalp chela manus and generally pronounced carination of the manus, especially the markedly granular rlm and rlvs carinae (figs. 17C, D, 18A, B), and metasoma, but can be separated from the latter by their considerably larger size and by the clustered instead of moniliform granules of the pedipalp carinae.
INCLUDED SPECIES:
Kochius barbatus
(Williams, 1971)
,
stat. nov.
;
Kochius bruneus
(Williams, 1970)
;
Kochius cerralvensis
(Williams, 1971)
,
stat. nov.
;
Kochius insularis
(Williams, 1971)
;
Kochius hirsuticauda
(Banks, 1910)
;
Kochius magdalensis
(Williams, 1971)
;
Kochius punctipalpi
(Wood, 1863)
;
Kochius sonorae
(Williams, 1971)
;
Kochius villosus
(Williams, 1971)
,
stat. nov.
DISTRIBUTION:
Kochius
is endemic to
Mexico
(recorded from
Baja California
,
Baja California Sur
, and
Sonora
) and the
United States
(recorded from
Arizona
,
California
, and
Nevada
; fig. 5). All except
K. hirsuticauda
and
K. sonorae
, occurring in the Mojave and Sonoran (including
Colorado
) deserts on the North American mainland, are endemic to the Baja
California
Peninsula and adjacent islands.
NATURAL HISTORY: Species of
Kochius
inhabit desert to semidesert habitats from sea level to
1350 m
altitude. They have been collected under stones or other available debris on the ground, and on the surface at night, with UV light detection. Although
Sissom (2000)
suggested that these scorpions are burrowers, the absence of fossorial adaptations suggests otherwise. The cheliceral fingers are not enlarged, the pedipalp chela fingers are fairly slender, the tarsi are not laterally compressed and lack setal combs, and the metasoma lacks enlarged carinal processes and lobes. The habitat and habitus of
Kochius
are consistent with the lapidicolous ecomorphotype (
Prendini, 2001a
).
REMARKS: As redefined here, this genus accommodates species previously assigned only to Williams’ (1971a)
punctipalpi
group of
Vaejovis
, for which
Soleglad and Fet (2008)
devised the name
Kochius
, without quantitatively testing its monophyly or composition.
Kochius
, as defined by
Soleglad and Fet (2008)
, was consistently polyphyletic, and the group of species hereby assigned to it consistently monophyletic, in the phylogenetic analyses of González-Santillán and Prendini (in press) based on morphology and those based on morphology and DNA. The following species are therefore transferred to other genera in the present contribution:
Mesomexovis atenango
,
comb. nov.
, previously assigned to the
punctipalpi
group (
Francke and González-Santillán, 2007
) and then to
Kochius
(
Soleglad and Fet, 2008
)
;
C. cazieri
,
comb. nov.
,
C. crassimanus
,
comb. nov.
, and
C. russelli
,
comb. nov.
, previously assigned to the
punctipalpi
group (
Williams, 1971a
, b;
Sissom, 2000
) and then to
Kochius
(
Soleglad and Fet, 2008
)
;
C. kovariki
,
comb. nov.
, previously assigned to
Kochius
(
Soleglad and Fet, 2008
)
.
The analyses of González-Santillán and Prendini (in press) identified sufficient, consistent diagnostic character differences to elevate to the rank of species,
K. barbatus
,
stat. nov.
, and
K. cerralvensis
,
stat. nov.
, previously considered subspecies of
K. punctipalpi
, and
K. villosus
,
stat. nov.
, previously considered a subspecies of
K. bruneus
(
Williams, 1971a
,
1980
;
Sissom, 2000
).
Vaejovis bruneus loretoensis
Williams, 1971
, was not found to differ consistently from
K. villosus
,
stat. nov.
, with which it is hereby synonymized:
Vaejovis bruneus loretoensis
Williams, 1971
=
Kochius villosus
(Williams, 1971)
,
syn. nov.
MATERIAL EXAMINED:
Kochius barbatus
(Williams, 1971)
,
stat. nov.
:
MEXICO
:
Baja California Sur
:
Municipio de La Paz
:
Las Cruces
,
25 ft
,
29.vii.1968
,
S.C. Williams
and
M.A. Cazier
,
holotype
♀
of
Vaejovis punctipalpi barbatus
Williams
, 1971
(
CAS
Type
No. 10469),
paratype
♀
(
AMNH
);
Las Cruces
,
4 km
NW
,
24
°
13.5108
′
N
110
°
7.41
′
W
,
31 m
,
24.vi.2008
,
H. Montaño
and
E. González
, 18 (
AMNH
[
ARA 2884
]),
18 (
IBUNAM
[
ARA 2274
]).
Kochius bruneus
(Williams, 1970)
:
MEXICO
:
Baja California Sur
:
Municipio de Comondú
:
San José de Comondú
,
5 mi.
SW
,
1000 ft
,
2.vii.1968
,
S.C. Williams
and
M.A. Cazier
,
holotype
♂
(
CAS
Type
No. 10410);
San José de Comondú
,
30 km
N
, between
Rosarito and San Isidro
,
26
°
21.2520
′
N
111
°
50.10
′
W
,
258 m
,
27.vi.2008
,
H. Montaño
and
E. González
, 28,
2♀
(
AMNH
[
ARA 2854
]).
Kochius cerralvensis
(Williams, 1971)
,
stat. nov.
:
MEXICO
:
Baja California Sur
:
Municipio de La Paz
:
Isla
Cerralvo
,
Bahía Limona
,
31.v.1969
,
S.C. Williams
,
holotype
♂
(
CAS
Type
No. 11470);
Isla
Cerralvo
,
Piedras Gordas
,
17.v.1970
,
S.C. Williams
and
V
.
F. Lee
, 18,
1♀
paratypes
(
AMNH
);
Isla
Cerralvo
, SW side,
24
°
10.9332
′
N
109
°
53.2332
′
W
,
50 m
,
27.v.2008
,
E. González
and
I.G. Nieto
, 18,
1♀
(
AMNH
[
ARA 2880
]).
Kochius hirsuticauda
(Banks, 1910)
:
U.S.A.
:
Arizona
:
Yuma Co.
:
Palm Canyon
,
Kofa Mountains
,
12.x.1968
,
M.A. Cazier
et al., 18,
1♀
(
AMNH
).
California:
San Diego Co.
:
Anza-Borrego Desert State Park
:
Pinyon Mountain Road
, ca.
4.8 km
towards
Pinyon Mountain Valley
from junction with
Route
S2,
33
°
03.043
′
N 116
°
23.270
′
W
,
870 m
,
30.viii.2005
,
R
.
Mercurio and L. Prendini
, 18,
1♀
(
AMNH
).
Kochius insularis
(Williams, 1970)
:
MEXICO
:
Baja California Sur
:
Municipio de La Paz
:
Isla
Partida
,
Central Valley
,
10.vii.1968
,
S.C. Williams
,
M. Bentzien
, and
B. Fox
,
holotype
♂
(
CAS
Type
No. 10422),
allotype
♀
(
CAS
).
Municipio de La Paz
:
Isla
Espíritu Santo
,
Playa Bonanza
,
24
°
27.3636
′
N
110
°
18.47
′
W
,
50 m
,
31.v.2008
,
E. González
and
I.G. Nieto
, 18,
1♀
(
AMNH
[
ARA 2875
]),
18,
1♀
(
IBUNAM
[
ARA 2260
]).
Kochius magdalensis
(Williams, 1971)
:
MEXICO
:
Baja California Sur
:
Municipio de La Paz
:
La Paz
,
75 mi.
NW
,
200 ft
,
4.vii.1968
,
S.C. Williams
and
M.A. Cazier
,
holotype
♂
(
CAS
Type
No. 10468);
El Pilar
, ca.
20 km
NE
Las Pocitas
,
24
°
28.7616
′
N
111
°
1.12
′
W
,
92 m
,
25.vi.2008
,
H. Montaño
and
E. González
, 18,
1♀
(
AMNH
[
ARA 2874
]),
18,
1♀
(
IBUNAM
[
ARA 2259
]);
Junction
Route
1 and road to
Juan de la Costa
,
5 km
N
,
24
°
08.500
′
N
110
°
28.566
′
W
,
30 m
,
9.vi.2005
,
E. González
,
W.E. Savary
,
R
.
Mercurio, and L. Prendini
, 18 (
AMNH
).
Kochius punctipalpi
(Wood, 1863)
:
MEXICO
:
Baja California Sur
:
Municipio de Los Cabos
:
Santiago
,
23
°
26.4
′
N 109
°
43.5666
′
W
,
225 m
,
9.vii.2004
,
O.F. Francke
,
E. González
, and
A. Valdez
, 18,
1♀
(
AMNH
[
ARA 1471
]);
San José del Cabo
, ca.
10 km
S
off
Route
1,
23
°
1.7622
′
N
109
°
43.49
′
W
,
50 m
,
10.vii.2005
,
W.E. Savary
,
E. González
, and
R
.
Mercurio
, 18,
1♀
(
AMNH
[
ARA 1473
]).
Municipio de La Paz
:
La Paz
,
15 km
SE
,
24
°
3.7332
′
N
110
°
10.53
′
W
,
476 m
,
8.vii.2004
,
O.F. Francke
,
E. González
, and
A. Valdez
, 18 (
AMNH
[
ARA 1472
]).
Kochius sonorae
(Williams, 1971)
:
MEXICO
:
Sonora
:
Municipio de Alamos
:
Alamos
,
11.2 mi.
W
,
23.i.1966
,
R
.
Winokur, C. Mays
, and
M.A. Nikerson
,
holotype
♂
(
CAS
Type
No. 10472).
Municipio de Navojoa
:
Microwave antenna Cerro Prieto
,
15 km
E
Navojoa
,
27
°
4.9830
′
N
109
°
17.4498
′
W
,
375 m
,
23.viii.2009
,
E. González
and
J.L. Castelo
, 18,
1♀
(
AMNH
[
ARA 3157
]);
Navojoa
,
32 km
S
,
12.ix.1966
,
J.
and
W. Ivie
, 18,
1♀
(
AMNH
).
Municipio de Soyopa
: Sierra el Encinal,
9 km
from Hwy
MEX 16
on dirt road to
El Encinal
,
28
°
35.4120
′
N 109
°
27.1476
′
W
,
380 m
,
24.vi. 2006
,
E. González
and
P. Berea
, 28 (
AMNH
[
ARA 2539
]);
dirt road between
El Encinal
and
Tonichi
,
28
°
33.2388
′
N
109
°
21.64
′
W
,
645 m
,
1.vii.2006
,
E. González
and
P. Berea
, 28 (
AMNH
[
ARA 3015
]).
Kochius villosus
(Williams, 1971)
,
stat. nov.
:
MEXICO
:
Baja California Sur
:
Municipio de Comondú
:
San Miguel Comondú
,
5–10 mi.
SW
,
800–1000 ft
,
3.vii.1968
,
S.C. Williams
and
M.A. Cazier
,
holotype
♂
of
Vaejovis bruneus villosus
Williams
, 1971
(
CAS
Type
No. 10467), 18,
1♀
paratypes
(
AMNH
).
Municipio de La Paz
:
San Evaristo
,
10–15 km
S
, dirt road,
24
°
47.7570
′
N
110
°
40.5018
′
W
,
50 m
,
12.vii.2008
,
H. Montaño
and
E. González
, 18 (
IBUNAM
[
ARA 2275
]),
18,
1♀
(
AMNH
[
ARA 2888
]).
Municipio de Loreto
:
Juncalito
,
25
°
49.8918
′
N
111
°
19.6908
′
W
,
5 m
,
20.v.2004
,
R
.
Mercurio
and
M. Nishiguchi
, 18,
1♀
(
AMNH
[
ARA 3132
]).
Municipio de Mulegé
:
San Ignacio
,
152 m
,
24.vi.1968
,
S.C. Williams
and
M.A. Cazier
,
holotype
♂
of
V
. b. loretoensis
(
CAS
Type
No. 10466),
allotype
♀
, 18,
1♀
paratypes
(
AMNH
).