Revision of the Central Asian scorpion genus Anomalobuthus Kraepelin, 1900, with descriptions of three new species and a generic synonymy (Scorpiones: Buthidae)
Author
Teruel, Rolando
Author
Kovařík, František
Author
Fet, Victor
text
Euscorpius
2018
270
1
45
journal article
1536-9307
3D3E9705-CEDD-4DBE-A23F-350C6DBE7972
Anomalobuthus rickmersi
Kraepelin, 1900
Figures 1–33
,
117–119
,
126–127
,
137
,
141
;
Tables 1
,
4–5
Anomalobuthus rickmersi
Kraepelin, 1900: 1–11
, figs. 4–7; Birula, 1904: 32; Birula, 1905: 449 (in part); Birula, 1911a: 73–74 Birula, 1911b: 171 (in part); Rickmer Rickmers, 1913: 442; Birula, 1917: 112 (in part);
Pavlovsky, 1934: 201
(in part).
Werner, 1934: 272
; fig. 341;
Weidner, 1959: 98
;
Stahnke, 1972: 122
(in part), fig. 3;
Vachon, 1974: 908
, fig. 37; Fet, 1980: 224 (in part); Fet, 1989: 80 (in part);
Sissom, 1990: 37
, 46, 48– 51, 96, 101, 120, figs. 3.17h, m; Fet, 1994: 531 (in part); Gromov & Kopdykbaev, 1994: 20 (in part); Fet et al., 1998: 613 (in part), fig. 6 [specimen from Repetek]; Fet & Lowe, 2000: 75 (in part, includes full list of references before 1998); Fet et al., 2001: 183–185, tab. 1 (in part). Fet et al., 2003: 4 (in part); Fet et al., 2005: 2– 3, 6–7, 10–11, 13, 22, 24–25, figs. 10, 23–25, tab. 1 (in part);
Graham et al., 2012: 95–96
(in part);
Warburg, 2013: 96
, 102 (in part);
Teruel et al., 2014: 1
, 4, 6–7, 9, fig. 20, tabs. 1–2 (in part).
Nec
Lourenço, 2001: 15–20
[misidentification, see below in Remarks section].
HOLOTYPE
♂
(
ZMUH
, examined).
Tajikistan
,
Khatlon Province
,
Kyzylsu River valley
, near Baljuvon (= Bal'dzhuvon or Baljuan), early summer 1896, leg.
W. Rickmer Rickmers. See
discussion below, in Remarks section.
OTHER MATERIAL EXAMINED
.
Turkmenistan
,
Karakum Desert
,
Lebap
(=
Chardzhou
)
Province
,
Repetek Nature Reserve
,
38°33'54"N
63°10'51"E
,
201 m
a.s.l.
,
21 July 1985
, leg
.
V
.
Fet
,
2♂♂
(
NHMW
),
5♂♂
,
1 juvenile
(
VFPC
); same data except
29 April 1967
, leg
.
V
.
I. Kuznetsov
,
1♂
(
ZSRO
Sc-1249); same data except
15 April 2002
, leg
.
V
.
Fet
,
1 juvenile
(
VFPC
);
Karakum Desert
,
Akhal Province
,
Tejen
(=
Tedzhen
,
Tedschen
),
37°22'39"N
60°29'52"E
,
188 m
a.s.l.
,
10–24 July 1904
[23 July–6 August], leg.
K. Aris
,
1♂
(
ZISP 809
);
Mary Province
,
Serkhetabad District
,
Badghyz Plateau
,
Chainury
,
35°40'52"N
62°01'34"E
,
458 m
a.s.l.
,
6 April 2002
, leg
.
V
.
Fet
& A
.
V
.
Gromov
,
1♀
(
FKCP
),
1♀
(
NMPC
)
.
Figures 1–3:
Anomalobuthus rickmersi
, adult male holotype: full-body views, dorsal (1) and ventral (2), plus original labels (3). Scale bar = 10 mm (1–2). The red inset at bottom right shows the label of the specimen examined by
Lourenço (2001)
, demonstrating it was not the actual Rickmers's type specimen.
DIAGNOSIS (emended). Adult size
30–34 mm
in males,
32–37 mm
in females. Coloration light yellow, essentially immaculate, only with some regular blackish spots on pedipalps and metasoma (darker and more extended in female); metasomal segment V and telson reddish brown (male) to black (female). Pedipalp fingers with 9–10 principal rows of denticles and 7–9 internal accessory denticles. Pectines with 23–27 teeth in males and
20–23 in
females. Tibial spur variable from absent (specimens from
Turkmenistan
) to present but very small on leg IV only (male
holotype
from
Tajikistan
). Metasoma conspicuously attenuate, with most carinae obsolete to vestigial; ventral lateral carinae of segment V composed of denticles narrowly conical (male) to narrowly lobate (female), dorsal lateral and lateral supramedian carinae of segments I–IV with terminal denticle enlarged; intercarinal spaces essentially smooth on segments I–III, sparsely granulose on ventral and lateral areas of IV and very densely granulose on ventral and lateral areas of V (granulation sparser in female). Telson vesicle elongate oval and sparsely setose.
Figures 4–7:
Anomalobuthus rickmersi
from Turkmenistan: adult male from Repetek (4–5) and adult female from Chainury (6– 7), full-body views, dorsal (4, 6) and ventral (5, 7). Scale bar = 10 mm.
Figures 8–13:
Anomalobuthus rickmersi
.
Figures 8–9.
Juvenile male from Repetek, Turkmenistan: full-body views, dorsal (8) and ventral (9). Scale bar = 10 mm.
Figures 10–13.
Adult male holotype: chelicerae, carapace and tergites I–III, dorsal (10), sternopectinal region and sternites III–V, ventral (11), distal part of left leg III, internal (12), distal part of left leg IV, internal (13).
REDESCRIPTION (adult male
holotype
).
Coloration
(
Figs. 1–2
) moderately faded and translucent due to long preservation, but still reliably traceable. Base light yellowish; in general, the base color is paler on pedipalp chelae, legs, and pectines. Chelicerae immaculate, except for dark brown finger teeth. Pedipalp femur and patella with internal surface infuscate; chela immaculate, only with finger denticles dark brown. Carapace immaculate, only with a blackish spot under every ocular group. Tergites immaculate. Coxosternal region and genital operculum immaculate. Pectines pale yellowish, immaculate. Sternites immaculate. Legs essentially immaculate; claws with distal half dark brown. Metasoma not conspicuously bicolor, only becoming progressively darker and redder distally, segments I–IV with a diffusely annulated appearance (basal and distal parts of each segment faintly infuscate as thin brown rings, which become larger and more diffuse dorsally) and segment V faintly infuscate, becoming progressively darker and redder distally; carinae not infuscate nor underlined with dark pigment. Telson vesicle reddish brown, aculeus with basal half yellowish and distal half dark brown.
Chelicerae
(
Fig. 10
). With dentition typical for the genus, as described for
A. krivochatskyi
sp. n.
(see below).
Pedipalps
(
Figs. 14–22
). Relatively short but very slender, essentially bare. Femur subtly curved inwards, with carinae weak, granulose to subdenticulate; intercarinal tegument smooth and glossy. Patella straight, with carinae obsolete to absent, smooth except on internal surface where vestigially denticulate; intercarinal tegument smooth and glossy. Chela elongate and very slender; manus conspicuously narrower than patella (ratio 0.79), subcylindrical (1.80 times longer than wide, 1.07 times wider than deep), with carinae weak to moderate, smooth; intercarinal tegument smooth and glossy; fingers very long (movable finger 2.44 times longer than underhand), only subtly curved and with 10 principal rows of denticles (the two basalmost rows are moderately well-defined), basal lobe/notch combination absent, external accessory denticles absent, internal accessory denticles very large and claw-like (increasing in size distally), numbering eight on both fixed and movable fingers, movable finger with one claw-like accessory denticle basal to the very large terminal denticle.
Carapace
(
Fig. 10
). Very strongly trapezoidal (much narrower anteriorly) and wider than long; anterior margin shallowly convex, with 5–6 pairs of thin macrosetae and some very short microsetae. Carination essentially absent: the only carinae present are the superciliaries, which are smooth. Furrows: anterior median, median ocular, central median, posterior median and posterior marginal fused, wide and moderately deep; lateral oculars, lateral centrals, central transverse, and posterior laterals long, narrow and shallow. Tegument very finely and densely granulose, with a few small to medium-sized granules scattered all over.
Sternum
(
Fig. 11
). Standard for the genus, relatively small and widely triangular in shape.
Genital operculum
(
Fig. 11
). Slightly damaged. Relatively large, each half roundly subtriangular in shape. Genital papillae present.
Pectines
(
Fig. 11
). Standard-sized for the genus: very long, extending beyond leg IV coxa-trochanter joint), subrectangular and densely setose. Tooth count 25/25. Basal plate heavily sclerotized, much wider than long, anterior margin with a very deep, narrow anteromedian furrow, posterior margin very shallowly convex.
Legs
(
Figs. 12–13
). Very slender, with all carinae weak and subgranulose to subcostate; intercarinal tegument smooth and glossy. Tibial spurs entirely absent on legs III, reduced to a sinuous remnant on legs IV.
Mesosoma
(
Figs. 10–11
). Tergites with the same sculpture as on carapace; I–VI irregularly tricarinate: the median longitudinal carina is weak, short, and formed by irregular medium-sized granules that do not project beyond posterior margin, but the submedian carinae are defined only by few coarse granules irregularly aligned; tergite VII with five well-defined carinae: the median carina is very short but strongly subserrate, the submedians and laterals are long, strong and finely serrate to denticulate. Sternites sparsely setose; III–VI glossy and with very subtle vestiges of smooth submedian carinae, spiracles relatively short and slit-like, almost transversely arranged (not strongly oblique), V with smooth patch absent; VII with two pairs of carinae: the submedians are long and finely crenulate to subcostate and the laterals are very short and subcrenulate, intercarinal tegument coriaceous to minutely granulose.
Metasoma
(
Figs. 23–25
). Elongated and slightly wider subdistally; with 10/10/10/8/5 complete to almost complete carinae, almost all obsolete to vestigial and smooth, with coarse punctations scattered: dorsal laterals very weak on I–III (with one terminal denticle enlarged), obsolete to vestigial as rounded ridges on IV, absent on V; lateral supramedians very weak on I–III (with one terminal denticle enlarged), obsolete to vestigial as rounded ridges on IV–V; lateral inframedians obsolete on I–III, absent on IV–V; ventral laterals obsolete on I– IV, weak to moderate on V, where become progressively stronger and somewhat flared distally, formed by sharply serrate, subequal denticles; ventral submedians obsolete on I–IV, absent on V; ventral median absent on I–IV, weak to moderate on V. Intercarinal tegument smooth and glossy, with minute but sharp granulation on lateral and ventral surfaces, which become much denser towards distal segments. Dorsal furrow obsolete on all segments. Setation sparse, mostly represented by 5–8 dark macrosetae over every carina.
Telson
(
Figs. 26
). Sparsely setose, with a few setae scattered on dorsal and lateral surfaces. Vesicle elongate oval (2.50 times longer than wide, 0.95 times wider than deep), tegument smooth and glossy, with vestiges of coarse granules arranged into three obsolete longitudinal carinae (ventral median and ventral submedians), and ventrally with some coarse punctations. Subaculear tubercle absent. Aculeus conspicuously shorter than vesicle, thick and shallowly curved.
Figures 14–26:
Anomalobuthus rickmersi
, adult male holotype: right pedipalp trochanter and femur, dorsal (14) and ventral (15), right pedipalp patella, dorsal (16), external (17) and ventral (18), right pedipalp chela, dorsal (19–20), external (21) and ventrointernal (22), metasoma and telson, lateral (23), dorsal (24) and ventral (25), telson, lateral (26).
FEMALE (Chainury,
Turkmenistan
). Very similar to male, sexual dimorphism evident by:
1)
size comparatively larger inside each size-class;
2)
coloration darker and much more sharply patterned, e.g., pedipalp femur with a conspicuous blackish spot on dorsodistal apex, which diffusely continues as a broad stripe over almost all internal surface, patella with internal surface blackish, carapace with anterior margin deeply infuscate, metasomal segment V and telson black;
3)
mesosoma
Teruel, Kova
ř
ík & Fet: Revision of Genus
Anomalobuthus
9
Figures 27–33:
Anomalobuthus rickmersi
from Turkmenistan, adult male from Repetek (27–30) and adult female from Chainury (31–33): metasoma and telson, lateral (27, 31), dorsal (28, 32) and ventral (29, 33), telson, lateral (30).
and metasoma slightly less slender;
4)
genital papillae absent;
5)
pectines smaller, with consistently lower tooth counts;
6)
metasomal segment V with lateral intercarinal tegument less granulose. See
Figs. 6–7
,
31–33
and
Tabs. 1
,
4–5.
VARIATION. Adult size varied from
30.4–34.1 mm
in males and
32.1–36.5 mm
in females.
Pectinal tooth count varied as follows: 23–27 teeth in males and
20–23 in
females, without clearly defined mode in either sex (
Tab. 5
).
Males
from
Repetek
are virtually identical to the
holotype
in all diagnostic characters.
The
only exception is that the tibial spurs are usually entirely absent.
This
character alone is inappropriate to separate both populations taxonomically, especially when only a single specimen from
Tajikistan
is available.
In
addition, the male from
Tejen
(
ZISP 809
) has tibial spurs on both legs
IV
as noted already by Birula (1905: 450)
COMPARISON. Adults of
A. rickmersi
can be very easily distinguished by the greater slenderness of the metasoma, especially in males (
Figs. 1–2
,
4–7
,
117
;
Tabs. 1
,
4
); even to unaided eye, most other congeners are more robust (
Figs. 34–45
,
65–66
,
79–82
,
90–91
,
107, 110, 113–114
;
Tabs. 2–4
). For example, see the following male ratios: metasoma + telson length / carapace length = 7.76
vs.
6.26–7.14, metasomal segments length / width = 1.74
vs.
1.02–1.35 (I), 2.13
vs.
1.35–1.82 (II), 2.20
vs.
1.39–1.88 (III), 2.56
vs.
1.75–2.23 (IV) and 2.71
vs.
1.85–2.38 (V).
Moreover, this species is the only one that has metasomal segment V and telson light reddish-colored in adult males (
Figs. 1–2
,
4–5
,
23–25
,
27–29
,
117
); in all other
Anomalobuthus
spp.
, it is blackish, entirely or at least in its distal half (
Figs. 34–35
,
38–39
,
59–61
,
79–80
,
84–86
).
DISTRIBUTION (
Fig. 141
). Widespread in southern Central Asia, from the sands of the Karakum Desert (eastern
Turkmenistan
), to the loess soils of the Kyzylsu River valley (southwestern
Tajikistan
). See further details below, in Remarks section.
ECOLOGY. According to the personal data of the collector (Rickmer Rickmers, 1913: 442), the
holotype
was collected in early summer, from crevices or cracks of fine alluvial soil (“
loess crannies
”). Populations from the Karakum Desert inhabit sand dunes but tend not to occupy loose sands where two other sympatric ultrapsammophiles are found,
Liobuthus kessleri
and
Pectinibuthus birulai
Fet, 1984
. Detailed ecological information from Repetek was published by Fet (1980).
NOTES. The
holotype
of
A. rickmersi
remains the single specimen known from
Tajikistan
. Its collecting site was reported by
Kraepelin (1900)
as Bukhara (“Bucharei”, see the original museum label in
Fig. 3
) and interpreted by all subsequent authors as originating from the city, the capital of the homonymous Bukhara Khanate, located in modern
Uzbekistan
. In addition, Birula (1911b: 171) erroneously stated that the type specimen of
A. rickmersi
originated from “westlichen Buchara” (i.e. western part of the Bukhara Emirate).
This
specimen was collected by the famous
German mountain
explorer
Willi Rickmer Rickmers
(1873–1965) who repeatedly visited the
Bukhara Emirate
, then a protectorate of the Russian Empire, which included part of modern
Uzbekistan and Tajikistan
(Rickmer-Rickmers, 1899; Rickmer-Rickmers, 1913).
During
preparation of the present revision, we discovered that
Rickmer Rickmers
(1913: 442) himself actually published unequivocal information on the precise collection locality and date of the
holotype
, which we reproduce below
verbatim
:
“On the way to Baljuan I noticed that many poplar trees were pitted with wormholes bored by that stately Capricorn beetle the
Pachydissus Sartus
. In
the early summer of 1896 the loess crannies around had yielded up to my collection new specimens of poisonous spider and scorpion,
Gylippus Rickmersi
and
Anomalobuthus Rickmersi
. I
feel highly gratified that such vicious creatures should bear my name.”
By the way, the “poisonous spider” mentioned by the impressed collector is a solpugid, also described by
Kraepelin (1899)
. It is a member of the
Gylippidae
family, currently regarded as a junior synonym of
Gylippus ferganensis
Birula, 1893
(see Harms & Dupérré, 2018). As all solpugids, it is indeed “vicious” but not “poisonous”.
Baljuan (also spelled Baldzhuan, now Baljuvon or Bal'dzhuvon) is a well-known town in modern
Tajikistan
, in the valley of Kyzylsu River, a tributary of the larger Panj River (= Pyandzh, Piandj), itself a tributary of the great Amudarya (the ancient Oxus). Baljuvon stands on the Silk Road, not far from the ancient
Bactra
(now
Balkh
in northern
Afghanistan
), and was last time noted as the place where Enver Pasha, a rival of the Turkish leader Mustafa Kemal (Ataturk), was killed in
1922 in
a skirmish with the Soviet troops.
In more peaceful times, Rickmer-Rickmers (1899) published a map of his fourth travel to the Bukhara Emirate, conducted in 1898 (the 1896 trip was his third travel). We can see that he followed the same and (then) only route from
Dushanbe
(now the capital of
Tajikistan
) to Khovaling, via Baljuvon. Baljuvon lies very far to the southeast from the former Emirate's capital city of Bukhara (modern
Uzbekistan
) and is isolated by several mountain ranges from the lowland deserts of
Uzbekistan
.
Although quite active zoological research was conducted in Central Asia by numerous Russian (and then Soviet) researchers, very few scorpions have been collected in the remote valleys of northern tributaries of Amu Darya in modern
Uzbekistan
and
Tajikistan
. It is not surprising, therefore, that no other specimens of
A. rickmersi
have been collected in this area since 1896; in fact, no other scorpion collections we know of originate from Baljuvon. Moreover, another, much more amazing scorpion from the same area was not discovered until the 1990s: the relict
Pseudochactas ovchinnikovi
Gromov, 1998
(Pseudochactidae)
. Its
type
locality lies in the Surkhandarya River valley, which separates modern
Uzbekistan
from
Tajikistan
(
Gromov, 1998
; Fet et al., 2004).
The
first known specimen from
Turkmenistan
(then the
Transcaspian Region
of the Russian Empire) was reported by
Birula
(1904: 32, 35).
It
was a female collected next to the railway station of
Repetek
in the
East Karakum
, the future site of the famed
Repetek Natural Reserve
, by the coleopterologist
Eduard Nikolaevich Fischer
in
February 1904
(
ZISP 808
).
The
second specimen was also published by
Birula
(1905: 449–450); it was a male from
Tejen
(=
Tedzhen
,
Tedshen
) in
Akhal Province
, collected by
K. Aris
in
June 1904
(
ZISP 809
).
Further
collections from
Turkmenistan
published by
Fet
(1989), included a series of more than
20 specimens
from
Repetek
(
ZISP
,
ZMMSU
). In addition,
two males
from Repetek are housed in
NHMW
and one in
R
. Kinzelbach's collection (
ZSRO
). Some of these specimens are illustrated here in
Figs. 4– 5
,
27–30
,
103–115. A
few specimens from northern
Turkmenistan
(Kunya-Urgench,
ZISP 1699
; Shakhsenem,
ZMMSU
Tb-374), published by Fet (1989) as
A. rickmersi
, actually correspond to a different species (
A. pavlovskyi
sp. n.
, see below)
.
|
Dimensions (mm)
|
TJK, Baljuvon
♂
holotype (ZMUH)
|
TKM, Chainury
♀
(FKCP)
♀
(FKCP)
|
| Carapace |
L / Wp |
3.05 / 3.80 |
3.67 / 4.30 |
3.55 / 4.03 |
| Mesosoma |
L |
7.70 |
9.40 |
7.10 |
| Tergite VII |
L / W |
2.15 / 3.50 |
2.35 / 4.10 |
1.79 / 4.18 |
| Metasoma + Telson |
L |
23.38 |
23.41 |
21.48 |
| Segment I |
L / W / D |
2.80 / 1.61 / 1.43 |
2.80 / 1.95 / 1.79 |
2.50 / 1.75 / 1.43 |
| Segment II |
L / W / D |
3.40 / 1.60 / 1.58 |
3.48 / 1.78 / 1.85 |
3.00 / 1.63 / 1.58 |
| Segment III |
L / W / D |
3.80 / 1.73 / 1.73 |
3.85 / 1.85 / 1.90 |
3.43 / 1.68 / 1.73 |
| Segment IV |
L / W / D |
4.43 / 1.73 / 1.74 |
4.35 / 1.88 / 1.95 |
4.05 / 1.75 / 1.74 |
| Segment V |
L / W / D |
4.55 / 1.68 / 1.50 |
4.78 / 1.97 / 1.72 |
4.45 / 1.78 / 1.50 |
| Telson |
L |
4.40 |
4.15 |
4.05 |
| Vesicle |
L / W / D |
2.63 / 1.05 / 1.10 |
2.65 / 1.40 / 1.30 |
2.60 / 1.30 / 1.20 |
| Aculeus |
L |
1.77 |
1.50 |
1.45 |
| Pedipalp |
L |
11.08 |
11.15 |
10.65 |
| Femur |
L / W |
3.03 / 0.65 |
2.95 / 0.75 |
2.75 / 0.70 |
| Patella |
L / W |
3.40 / 0.95 |
3.40 / 1.08 |
3.25 / 1.00 |
| Chela |
L |
4.65 |
4.80 |
4.65 |
| Manus |
L / W / D |
1.35 / 0.75 / 0.70 |
1.35 / 0.86 / 0.85 |
1.11 / 0.75 / 0.75 |
| Movable finger |
L |
3.30 |
3.45 |
3.54 |
| Total |
L |
34.13 |
36.48 |
32.13 |
Table 1:
Measurements of three adults of
Anomalobuthus rickmersi
.
Abbreviations: length (L), width (W), posterior width (Wp), depth (D).
It is clear from A. Birula's publications that he never studied the
holotype
of
A. rickmersi
; however, it was seen and its trichobothrial pattern sketched by
Vachon (1974: 908
, fig. 37). On the other hand,
Lourenço (2001)
claimed to have examined it; however, it is obvious that the specimen he received on loan from ZMUH was
not
Kraepelin's original
holotype
, but a misidentified and wrongly labeled juvenile of maybe even another genus.
Lourenço (2001)
declared this specimen as an immature with a total length of
13–14 mm
[
sic
], pectinal tooth count 24–26, and textually wrote: “...
cutting edge of the movable finger with 6 series of granules
[...]
the basalmost extending onto more than half the length of the finger.
[... Pectinal]
Fulcra almost vestigial and flat...
” [originally in French, English translation and bracketed text added by us].
None of these crucial characters match the
holotype
of
A. rickmersi
and clearly contradict the original description and illustrations published by
Kraepelin (1900)
, which were confirmed by our own examination of the true type: it is a large adult male with a total length of
34 mm
, pectinal tooth count 25/25, fingers with 10 rows of primary denticles of which the basal-most extends onto less than one-third of finger, and fulcra normally developed and pearl-like. Finally, a photo of the label of the specimen examined by
Lourenço (2001)
, provided by himself to V.F.in 2009, does not correspond to any of those accompanying the actual
holotype
(
Fig. 3
).