Revision of some historical types of the genus Lithobius Leach, 1814 (Chilopoda, Lithobiomorpha) from the 1949 / 50 Austrian Iran Expedition, with new molecular data for L. iranicus Attems, 1951
Author
Akkari, Nesrine
3 Zoological Department, Naturhistorisches Museum Wien, Burgring 7, 1010 Vienna, Austria.
Author
Macek, Oliver
0000-0002-8146-5373
3 Zoological Department, Naturhistorisches Museum Wien, Burgring 7, 1010 Vienna, Austria.
oliver.macek@nhm-wien.ac.at
Author
Edgecombe, Gregory D.
0000-0002-9591-8011
The Natural History Museum, Cromwell Road, London SW 7 5 BD, United Kingdom.
g.edgecombe@nhm.ac.uk
text
Zootaxa
2024
2024-03-15
5424
4
401
422
http://dx.doi.org/10.11646/zootaxa.5424.4.1
journal article
10.11646/zootaxa.5424.4.1
1175-5326
10821452
3BDADC13-421B-4F90-BFFE-E0C0158DCF9E
Lithobius
(
Lithobius
)
iranicus
Attems, 1951
Figs 1–5
,
6A–H, J
,
7
,
8A–C
,
9–10
Lithobius
(
Alokobius
)
iranicus
Attems, 1951: 403
, fig. 15.
Lithobius rhiknus
Attems, 1951: 403
, fig. 16.
Syn. nov.
Lithobius inaequidens
Attems, 1951: 404
, fig. 17.
Syn. nov.
Diagnosis.
Lithobius
(
Lithobius
)
with 44–52 antennal articles. Ocelli 1+12–1+23, in four rows; Tömösváry’s organ about as large as adjacent ocelli. Forcipular coxosternal margin most commonly with 4+4 teeth alternating in size from small to large distally, varying from 3+4 or 4+3 to 4+5 or 5+6, innermost tooth the most posterior; spiniform porodont distal to outermost tooth (4+4) or between outer pair of teeth (>4 teeth). Angulation or short projection on T9, well developed projections on TT11 and 13. Coxal pores usually ovate, slit-shaped in some large males. Female gonopod with 3+3 spurs, inner spur small; claw bipartite. Males with pair of dorsal sulci on femur and tibia of legs 14 and 15, on femur and variably tibia of leg 13. Legs 13–15 consistently with DaF and DpF; 13DaT, 13DpT, 14VaC, 15VaC consistently present.
Material examined.
Lectotype
:
IRAN
●
♀
NHMW
MY3963
; lectotype here designated; “Lahidschan, Buchenwald Löffler F106”.
Paralectotypes
:
●
2 ♂♂
NHMW
MY10383
,
10384
;
Lahidschan
, same collection as lectotype
●
1 ♂
NHMW
MY3964
; “Löffler H.57”
.
Non-type material:
Lithobius iranicus
:
IRAN
●
1 ♀
NHMW
MY5700
●
1 ♂
NHMW
MY5701
; “F101” [Lahidschan]
●
1 ♂
NHMW
MY10387
; close to a road
between Razekeh and Peren
;
36°17'42.86"N
,
52°21'47.26"E
,
424 m
;
15.12.2017
;
H. Ghanavi
leg.
Other material:
Lithobius inaequidens
Attems, 1951
.
IRAN
●
1 ♂
syntype
NHMW
MY3962
;
“F111”, “Tscheran Ham” ●
1 ♂
syntype
NHMW
MY10385
; “
Teheran
Hain I.50”
Lithobius rhiknus
Attems, 1951
.
IRAN
●
1 ♀
syntype
NHMW
MY3942
; “
Mazandaran
, H82 Löffler”; ●
1 ♂
syntype
NHMW
MY4044
; “
Mazandaran
, H87”.
Description based on female
lectotype
.
Size
. Length (anterior margin of cephalic plate to posterior margin of telson)
21 mm
(
Fig. 1A
).
Colour
(in ethanol). Cephalic plate orange-yellow; anterior tergites yellow, darker orange-yellow from posterior half of trunk; antennae and legs yellow (
Fig. 1A, B
).
Cephalic plate
. Width slightly greater than length. Lateral marginal interruption distinct. Posterior border faintly convex backwards (
Fig. 1B
).
Antennae
. 50/48 articles, extending back to anterior part of T7 (
Fig. 1A
). Ultimate article twice as long as penultimate.
Ocelli
. 1+16 and 1+13 ocelli (left and right side of cephalic plate, respectively), in four rows (4, 4, 3, 4). Posterior ocellus ovoid, its long axis nearly twice the diameter of largest seriate ocellus.
Tömösváry’s organ.
Slightly separated at anteroventral edge of ocellar field, about as large as adjacent ocelli (
Fig. 1D
).
Forcipular coxosternite
. 4+4 teeth; inner tooth the smallest, positioned posterior to the others; second and fourth (outer) teeth equal sized, third tooth distinctly smaller (
Figs 1C
,
2A
). Median diastema shallow, curved. Porodont at a small, blunt shoulder; porodont spiniform, tapering to a point, translucent (
Fig. 2 C
). Anterior part of coxosternite bearing numerous long setae.
Tergites
. T1 95% width of cephalic plate and T3. T9 with a triangular projection on left side and an angulation on right; TT11 and 13 with strong projections, slenderer on T13 (
Fig. 2B
).
Legs
. Tarsal articulation well defined on all legs. Leg 15 with accessory apical claw. Plectrotaxy as in
Table 2
.
Coxal pores
. 5565/5665, ovate, separated by less than their diameter (
Fig. 2D
).
Female gonopods
. 3+3 conical spurs, inner spur about 60% length of largest (outer) spur. Claw bipartite, its two denticles subequal in size or outer spur slightly larger. First article of gonopod bearing 10–12 ventral setae, second article nine, third article three (
Figs 2D
,
8A
).
FIGURE 1.
Lithobius iranicus
Attems, 1951
, ♀ lectotype NHMW MY3963.
A.
Habitus, dorsal view.
B.
Cephalic plate and T1, dorsal view.
C.
Forcipular segment, ventral view.
D.
Ocelli and Tömösváry’s organ, lateral view.
Abbreviation
:
TO
= Tömösváry’s organ.
FIGURE 2.
Lithobius iranicus
Attems, 1951
, ♀ lectotype NHMW MY3963.
A.
Close-up of coxosternal margin showing teeth and porodont, ventral view.
B.
TT9-13, dorsal view.
C.
Porodont and lateralmost teeth, ventral view.
D.
SS13-15 posterior segments, ventral view.
Abbreviation
:
Po
= porodont.
Description of the newly collected specimen (male NHMW MY10387).
Size
. Length (anterior margin of cephalic plate to posterior margin of telson)
24.5 mm
(
Fig. 3A
).
Colour
(in 96% ethanol). Cephalic plate purple-brownish-yellow; anterior tergites purple-yellow, darker from posterior half of trunk; antennae and legs pale purple-yellow (
Fig. 3A, B
).
Cephalic plate
. Width slightly greater than length. Lateral marginal interruption distinct. Posterior border faintly convex backwards (
Fig. 3B
).
Antennae
. 45+ (broken)/50(detached) articles (
Fig. 3A, B, D
).
FIGURE 3.
Lithobius iranicus
Attems, 1951
, ♂, new specimen NHMW MY10387.
A.
Habitus, dorsal view.
B.
Cephalic plate and T1, dorsal view.
C.
Ocelli, lateral view.
D.
Forcipular segment, ventral view.
Ocelli
. 1+16 and 1+15 ocelli (left and right side of cephalic plate, respectively), in four rows. Posterior ocellus ovoid, its long axis nearly twice the diameter of largest seriate ocellus.
Tömösváry’s organ.
Slightly separated at anteroventral edge of ocellar field, about as large as adjacent ocelli (
Fig. 3C
).
Forcipular coxosternite
. 4+4 teeth; inner tooth positioned posterior to the others; second and fourth (outer) teeth larger, equal sized; third tooth similar in size to inner one (
Figs 3D
,
4A
). Median diastema V-shaped. Porodont translucent, shorter and stouter than in the described
type
(
Fig. 4A
). Anterior part of coxosternite bearing numerous long setae.
Tergites
.T1 almost 95% width of cephalic plate and T3. T9 with short, blunt projections; TT11 and 13 with strong projections, slenderer on T13 (
Fig. 4B
).
Legs
. Tarsal articulation well defined on all legs. Leg 15 with accessory apical claw. Plectrotaxy as in
Table 7
.
Coxal pores
. 7776/7766, ovate, separated by less than their diameter (
Fig. 4C
).
FIGURE 4.
Lithobius iranicus
Attems, 1951
, ♂, new specimen NHMW MY10387.
A.
Close-up of coxosternal margin showing teeth and porodont.
B.
TT9-13, dorsal view.
C.
Posterior segments, ventral view.
Abbreviation
:
Po
= porodont.
Male sexual characters
. Legs 13 with paired sulci on prefemur,14 and 15 with paired sulci on prefemur and femur, faintly visible on tibiae (
Fig. 5B
). Posterior margin of first genital sternite gently concave. Gonopod twosegmented, both articles bearing several setae (
Fig. 4C
).
Variability.
See
Table 1
for meristic characters and
Tables 2–9
for plectrotaxy of all specimens referred to
L. iranicus
and its subjective synonyms.
L. inaequidens
syn. nov.
and
L. rhiknus
syn. nov.
Variably present spines are 13VpT (usually present), 14VmT (usually present), 14DaT (usually present), 15VmT (present or absent in similar frequency), and 15DpT (usually absent). Between
type
specimens, either second or fourth coxosternal tooth positioned most anteriorly. Median diastema ranging from shallow to relatively deeply U- or V-shaped.
Males with pair of dorsal sulci on (prefemur) femur and tibia of legs 14 and 15, on femur and variably tibia of leg 13 (
Fig. 5
).
Attems (1951)
gave similar type localities for his nominal species
Lithobius inaequidens
syn. nov.
: “Tscheran Ham. Lahidschan, Buchenwald. Sabzawaran”. The label data accompanying the specimens bear different codes, viz. F111, which is according to
Starmühlner & Edlauer (1957)
: Lahijan (
Gilan
), loamy steep slope (N-exposed). We chose the same coordinates as the
lectotype
of
L. iranicus
. The second label “
Teheran
Hain I.50”, which translates to “
Teheran
grove”, cannot be assigned to a precise locality. The locality “Sabzawaran” corresponds today to Jiroft with a precision of ca
20 km
. No label data associated with Attems’ material indicate what specimen(s) of
L. inaequidens
came from Sabzawaran and, given that is far to the south of the two records that can be corroborated, we do not map its occurrence in Sabzawaran/Jiroft.
TABLE 2.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, lectotype female (NHMW MY3963). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
1–2
|
- |
- |
mp |
mp |
m |
- |
- |
ap |
ap |
a |
3–11
|
- |
- |
mp |
amp |
amp |
- |
- |
ap |
ap |
ap |
12
|
- |
m |
amp |
amp |
amp |
- |
- |
amp |
ap |
ap |
13
|
- |
m |
amp |
amp |
amp |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
a |
a |
- |
amp |
ap |
p |
15
|
a |
m |
amp |
amp |
a |
a |
- |
amp |
ap |
- |
TABLE 3.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, paralectotype male (NHMW MY10383). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
13
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
a |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
a |
a |
- |
amp |
ap |
p |
TABLE 4.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, paralectotype male (NHMW MY10384). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
13
|
a |
m |
amp |
amp |
amp |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
- |
TABLE 5.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, paralectotype male (NHMW MY3964). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
13
|
- |
m |
amp |
amp |
amp |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
p |
TABLE 6.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, non-type male (NHMW MY5701). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
13
|
a |
m |
amp |
amp |
amp |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
- |
TABLE 7.
Plectrotaxy of
Lithobius iranicus
Attems, 1951
, new male specimen (NHMW MY10387). C—coxa; F— femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
13
|
a |
m |
amp |
amp |
amp |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
- |
TABLE 8.
Plectrotaxy of
Lithobius inaequidens
Attems, 1951
syn. nov.
, syntype male (NHMW MY3962). C— coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
1–2
|
- |
- |
mp |
mp |
m |
- |
- |
ap |
ap |
a |
3
|
mp |
amp |
am |
- |
- |
ap |
ap |
ap |
4–12
|
- |
- |
mp |
amp |
am(p) |
- |
- |
ap |
ap |
ap |
13
|
- |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
- |
TABLE 9.
Plectrotaxy of
Lithobius rhiknus
Attems, 1951
syn. nov.
, syntype female (NHMW MY3942). C—coxa; F—femur; P—prefemur; t—trochanter; T—tibia; a—anterior, m—median, p—posterior.
Leg pair
|
Ventral
|
Dorsal
|
C
|
t
|
P
|
F
|
T
|
C
|
t
|
P
|
F
|
T
|
1–3
|
- |
- |
mp |
mp |
m |
- |
- |
ap |
ap |
a |
4–10
|
- |
- |
mp |
amp |
amp |
- |
- |
ap |
ap |
a |
11
|
- |
- |
amp |
amp |
amp |
- |
- |
ap |
ap |
ap |
12
|
- |
- |
amp |
amp |
amp |
- |
- |
amp |
ap |
ap |
13
|
- |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
14
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
ap |
15
|
a |
m |
amp |
amp |
am |
a |
- |
amp |
ap |
- |
FIGURE 5.
Paired sulci on ultimate legs of males, dorsal view.
A–B.
Lithobius iranicus
Attems, 1951
.
A.
Paralectotype NHMW MY10383.
B.
NHMW MY10387.
C.
Lithobius rhiknus
Attems, 1951
syn. nov.
, leg 13, syntype NHMW MY4044.
D.
Lithobius inaequidens
Attems, 1951
syn. nov.
, NHMW MY3962. Arrows pointing to sulci.
We opted against the selection of a
lectotype
for the name-bearing types of
Lithobius rhiknus
syn. nov.
as it remains impossible to pinpoint the type localities with precision as according to the label data, both
syntypes
originate from two different localities in
Mazandaran
(“H82” and “H87”), a province also located along the southern coast of the Caspian Sea.
Attems (1951)
stated “Chorramabad. Dschungel bei Tschalous” (translating Khorramabad. Jungle in Chalus) as type localities of the species, which most likely also correspond to the cities Khorramabad and Chalus, both in Prov.
Mazandaran
.