4258 (2): 121 137 articlecase.asu.ru/files/form_312-27105.pdfand monotarsobius verhoeff, 1905 of the...

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Zootaxa 4258 (2): 121–137 http://www.mapress.com/j/zt/

Article

https://doi.org/10.11646/zootaxa.4258.2.2

http://zoobank.org/urn:lsid:zoobank.org:pub:949BAF3D-D8DF-4BC5-98CB-1F2D946EF802

Revision of Disphaerobius Attems, 1926 (Chilopoda: Lithobiomorpha:

Lithobiidae: Pterygoterginae), a centipede genus with

remarkable sexual dimorphism

GYULLI SH. FARZALIEVA1, PAVEL S. NEFEDIEV2 & IVAN H. TUF3,4

1Department of Zoology, Perm State University, Bukireva 16, Perm 614600, Russia. E-mail: [email protected] of Ecology, Biochemistry and Biotechnology, Altai State University, Lenina 61, Barnaul 656049, Russia. E-mail: [email protected] of Ecology and Environmental Science, Faculty of Science, Palacký University, Šlechtitelů 27, Olomouc 77900, Czech Republic. E-mail: [email protected] author. E-mail: [email protected]

Abstract

Disphaerobius Attems, 1926, the sole constituent genus of the subfamily Pterygoterginae, is synonymized with Ptery-

gotergum Verhoeff, 1934 syn. nov., and is shown to comprise only two species: D. svenhedini (Verhoeff, 1934) comb.

nov. ex Pterygotergum, and D. loricatus (Sseliwanoff, 1881) (= Lithobius magnificus Trotzina, 1895), both syn. nov. and

comb. nov. ex Lithobius Leach, 1814. Disphaerobius is rediagnosed, both of its basically Central Asian species are rede-

scribed from fresh material of both sexes, and their distributions are mapped. Disphaerobius svenhedini is newly recorded

from S Kazakhstan and W Mongolia. New material of D. loricatus is also recorded. Sexual dimorphic characters in Lith-

obiomorpha are briefly discussed, these being considered especially strongly pronounced and unique in Disphaerobius.

Key words: Disphaerobius, Pterygotergum, Lithobiidae, taxonomy, new synonymy, new combination, Kazakhstan, Mon-

golia, China

Introduction

Attems (1926, 1927), in his global keys to genera of the Lithobiinae, proposed the new genus Disphaerobius

Attems, 1926 to incorporate only a single species, Lithobius magnificus Trotzina, 1895. The latter species had been

described from a single male collected from the Mugodzhar Hills, southern spurs of the Ural Mountains, all lying

within NW Kazakhstan, Central Asia (Trotzina 1895). Unfortunately, the original description of L. magnificus was

not entirely adequate, partially even incorrect. Thus, Trotzina (1895) noted the presence of 6 teeth on the

coxosternite, instead of the correct 4 teeth + a pair of strong porodonts similar to teeth. This was exactly the

character that misled Attems (1926, 1927) to establish his Disphaerobius. Attems (1927) also mentioned certain

tergal modifications as being common to both sexes, although L. magnificus had been based on a single male: “14.

Tergit stark vergrößert, mit stark vorgewölbten Seiten, durch einen Eindruck in 2 flache Hemisphären geteilt.

Seitenränder des 8., 9., 10., 13–15. Tergits mit Höckerzähnen, 3+3 Kieferfußhüftzähne” [Tergite 14 greatly

enlarged with strongly bulging sides, resembling two flat hemispheres. Margins of tergites 8, 9, 10, 13–15 with

cusp teeth, 3+3 forcipular teeth]. To make it worse, Zalesskaja (1978), in her monograph of the Lithobiomorpha of

the former Soviet Union, totally omitted both the genus and species.

Lithobius loricatus Sseliwanoff, 1881 was first described, also based in a single male, but it came from the East

Kazakhstan Region of Kazakhstan (Sseliwanoff 1881). Farzalieva & Zalesskaja (2003) not only revised the

holotype, but also redescribed the species in due detail, based on new samples of both sexes. The material stemmed

from the steppe regions of Transuralia, Russia, an area adjacent to the Mugodzhar Hills, the terra typica of D.

magnificus. However, Farzalieva & Zalesskaja (2003) likewise omitted Trotzina’s (1895) paper on Lithobius

magnificus, the type-species of Disphaerobius.

Accepted by T. Wesener: 28 Feb. 2017; published: 27 Apr. 2017 121

Pterygotergum Verhoeff, 1934, another new monotypic genus, was originally described, based on

Pterygotergum svenhedini Verhoeff, 1934, from the Tian-Shan Mountains of NW China (Verhoeff 1934b). As the

type series contained material of both sexes, Verhoeff was so impressed by the degree of sexual dimorphism

observed both in the forcipular teeth and tergal structure that he created a new family, Pterygotergidae Verhoeff,

1934, to incorporate the genus and species alone. The diagnosis of the family reads as follows (with a few misprints

corrected here). “Der Trochanteropräfemur der Kieferfüsse ist verlängert (fig. 1, a), nach innen gebogen und der

Innenrand eingebuchtet, zugleich auffallend gebräunt und verdickt, das innere Ende der Trochanterkerbe liegt weit

hinter der Mitte und daher der inneren Endecke genähert. […] (fig. 3). Beim ♂ sind das 10. 12. und 14. Tergit

erweitert und zwar das 12. ganz kolossal und flügelartig. (fig. 5).” [Forcipular trochanteroprefemur is elongated

(fig. 1, a), bent inward and the inner margin concave, being strikingly tanned and thickened, the inner end of the

trochanter’s notch is located far behind the middle and therefore approaching the inner corner… (fig. 3). In the

male, the 10th, 12th and 14th tergites are expanded, in particular the 12th is really enormous and wing-shaped (fig.

5).] (Verhoeff 1934b: 28).

Since then, the Pterygotergidae has been downgraded and is currently treated as a subfamily, Pterygoterginae,

within the family Lithobiidae (Bonato et al. 2011). Pterygotergum is its sole component genus and has only

occasionally been considered in discussions on lithobiid classification and phylogeny (Eason 1992; Bonato et al.

2011), in checklists of the Chinese fauna (Wang & Mauriès 1996; Ma et al. 2014) or as a remarkable example of

sexual dimorphism in Chilopoda (Lewis 1981).

Prompted by new material coming from southern Kazakhstan and Mongolia, but especially given such a

unique synapomorphy as a laterally serrate and enlarged male tergite 14, all shared by the above three nominate

species, i.e. Disphaerobius loricatus (Sseliwanoff, 1881), D. magnificus (Trotzina, 1895) and Pterygotergum

svenhedini Verhoeff, 1934, we do not hesitate to formally synonymize both former species. Besides this,

Pterygotergum is to be synonymized under Disphaerobius. Thus, Disphaerobius becomes oligotypic, with D.

loricatus being quite widespread across the largely arid, steppe to semi-desert parts of Russia’s S Transuralia, as

well as W and E Kazakhstan, whereas D. svenhedini occurs in NW China, S Kazakhstan and W Mongolia (Fig.

30). In addition, a new, updated diagnosis of Disphaerobius is given, again the sole component genus of

Pterygoterginae.

Material and methods

We have re-examined the holotype of Lithobius loricatus, kept in the collection of the Zoological Institute of the

Russian Academy of Science, St. Petersburg (ZIL). We have also found and studied additional 48 specimens

coming from the steppe zone of the Orenburg Area, southern Urals, Russia, and the Aktobe Region of W

Kazakhstan. This additional material is shared between the holdings of the Zoological Museum of the Moscow

State University, Moscow (ZMUM) and the Perm State University, Perm, Russia (PSU).

Regrettably, the holotype of L. magnificus has not been relocated. As it is absent from the ZIL or ZMUM

collection, it seems safe to presume as being lost.

Only a single male syntype of Pterygotergum svenhedini is currently deposited in the Bavarian State

Collection of Zoology (Zoologische Staatssammlung München, ZSM), also revised here. The remaining syntypes

could have been expected to be housed in the Royal Museum of Natural History (Naturhistoriska riksmuseet) in

Stockholm, Sweden, but they are not there (Göran Andersson, in litt.). Additional five specimens of svenhedini

were collected by A.A. Fomichev from the Almaty Region, S Kazakhstan and from the Baitag Bogd Uul Mt.

Range, W Mongolia. These samples are currently deposited in the ZMUM and PSU.

The counts of tergites follows those accepted in Bonato et al. (2010).

Measurements. The total body length is measured from the fore margin of the cephalic plate to the rear edge of

the postpedal segment. Leg length excludes the claw. Lengths are presented as the minimum and maximum values

observed. All measurements are in millimeters (mm).

Plectrotaxy. Legs spinulation data are given in a tabular form. Variations in plectrotaxy within a single

specimen or between specimens are presented in parentheses. The number of coxal pores on legs 12–15 is given as

a formula where a sequence of Arabic numerals means the number of pores from legs 12 to 15.

FARZALIEVA ET AL.122 · Zootaxa 4258 (2) © 2017 Magnolia Press

Specimens were photographed using an Olympus Pen PM1 camera with an Olympus M.Zuiko Digital ED 60

mm f/2.8 Macro lens. Digital images were prepared using Photoshop CS6 image stacking software. SEM

micrographs were taken at the PSU with the help of a Hitachi TM3000 scanning electron microscope with a back-

scatter electron detector.

The following abbreviations are used in the text and tables: T/TT—tergite(s), S/SS—sternite(s), Tim—

intermediate tergite, V—ventral, D—dorsal, C—coxa, t—trochanter, P—prefemur, F—femur, Ti—tibia, Ts1—

tarsus 1, Ts2—tarsus 2, a—anterior, m—median, p—posterior.

Taxonomic part

Disphaerobius Attems, 1926

Disphaerobius—Attems, 1926: 382, 383.Disphaerobius—Attems, 1927: 243 (n. gen.), 244.Disphaerobius—Eason, 1992: 3, 5.Pterygotergum—Verhoeff, 1934b: 29 (gen. nov.) syn. nov.Pterygotergitum (sic!)—Lewis, 1981: 274, 428.Pterygotergum—Bonato et al., 2011: 388.

Type species: Lithobius loricatus Sseliwanoff, 1881

New diagnosis. Disphaerobius is characterized by the functionally biarticulated tarsi 1–13, the 20-segmented

antennae, the absence of posterolateral triangular projections on the tergites, the rounded edges of the

macrotergites, the 1-segmented male gonopods, the small Tömösváry’s organ (smaller than the nearest ocellus), the

average or large body size (18–41 mm), and the serrate and broadened tergites in the male (Table 1).

Description. Males are characterized by the following combination of secondary sexual features: serrate and

broadened TT 12–Tim (either T 13 or T 10) and a strongly modified ultimate femur showing a dorsal depression

with a rounded outgrowth distodorsally. Females with a simple gonopodal claw (a tiny tooth may be present on

claw’s external face); gonopods without true dorsolateral spines, but with rows of setae and with 2+2 or 3+3 spurs.

Forcipules extended significantly beyond cephalic capsule. Antennae with 20+20 segments. All tarsi functionally

biarticulated, with Ts1 about 2 times longer than Ts2. Tarsi 11–13 each with a ventral row of strong setae especially

well-developed on midbody legs.

Remarks. There are several taxa in Central Asia that share certain morphological characters with

Disphaerobius, e.g. Schizotergitius Verhoeff, 1930, species from the genus Hessebius Verhoeff, 1941 with

functionally biarticulated tarsi (H. perelae Zalesskaja, 1978, H. plumatus Zalesskaja, 1978, H. procurvus

Zalesskaja, 1978 and H. pervagatus Zalesskaja, 1978), as well as species of the giganteus-group of Lithobius,

subgenus Ezembius Chamberlin, 1919 as diagnosed by Eason (1986b) to accommodate several Central Asiatic

species. These taxa are characterized by the functionally biarticulated tarsi of legs 1–13, the antennae composed of

20 antennomeres, the rounded posterior angles of all tergites, the 1-segmented male gonopods, and Tömösváry’s

organ being equal in size to the nearest ocellus or smaller (Table 1). The other members of the subfamily

Lithobiinae present in the region differ from the above group of species in the following features: the genus

Australobius Chamberlin, 1920, which mainly occurs in Southeast Asia (Ma et al. 2008a, 2008b; Qin et al. 2014)

differs in the number of antennomeres (24–33, usually more than 25); the subgenera Sigibius Chamberlin, 1913

and Monotarsobius Verhoeff, 1905 of the genus Lithobius, as well as the genus Validifemur Ma, Song et Zhu, 2007

in the functionally monoarticulated tarsi 1–13.

Composition. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov., D. loricatus (Sseliwanoff, 1881) comb.

nov.

Distribution. A basically Central Asian distribution ranging from the Sol-Iletsk District in the Orenburg Area,

Russia in the west and north to the Xinjiang Uygur Autonomous Region, China and the Altai Mts in western

Mongolia in the east, and to the Zhambyl District in the Almaty Region, Kazakhstan in the south (Fig. 30).

Zootaxa 4258 (2) © 2017 Magnolia Press · 123DISPHAEROBIUS REVISION

TA

BL

E 1. D

iagnostic differences betw

een the genera D

isphaerobius, Schizotergitius, H

essebius and the giganteus-group of L

ithobius.

Disphaerobius A

ttem

s, 1926

giganteus-grou

p of Lithobius*

Schizotergitius V

erh

oeff, 1930

Cen

tral A

sian

sp

ecies of th

e gen

us

Hessebius V

erh

oeff, 1941**

Bod

y len

gth

20–41, usually 25–35 m

m

10–50 m

m

22–25 m

m

18–30 m

m

Nu

mb

er of ocelli

4–7

7–10

5–6

4–10, usually 5–7

Size of T

öm

ösváry’s

organ

very sm

all, m

ore than 2 tim

es sm

aller than

nearest ocellus

equal to nearest ocellus or sm

aller

sm

aller than nearest ocellus

sm

aller than nearest ocellus

Nu

mb

er of cox

ostern

al

teeth

2+

2 – 3+

3

2+

2

2+

2

2+

2

Porod

on

ts

thick and short, sim

ilar to adjacent teeth

variable: fro

m setiform

to thick and short

setiform

variable: fro

m setiform

to thick

and

short

Secon

dary sex

ual

ch

aracters in

m

ale T

T

serrate and broadened m

ight start fro

m T

10, alw

ays present on T

14 and T

im

absent (only in L

. giganteus m

acrotergites strongly

elongated)

absent

absent

Male leg 14

slightly thicker than previous legs, F

w

ith

tw

o dorsal sulci (absent in som

e

specim

ens)

variable: thickened, T

i w

ith distal expansion or

shallow

dorsal sulcus or not m

odified (only in

L

.

giganteus not thickened, F

and T

w

ith 1–2 dorsal

sulci (absent in som

e specim

ens))

not thickened, w

ithout visible

secondary sexual characters

slightly thicker than previous legs,

without visible secondary sexual

characters, rarely T

i and/or F

w

ith very

weak, hardly visible sulci

Male leg 15

thickened, F

w

ith tw

o distinct sulci, i.e. a

poorly expressed dorsolateral sulcus and a

deep sulcus, latter form

ing apically a

round tubercle

variable: thickened, T

i w

ith dorsal sulcus or not

modified (only in

L

. giganteus not thickened, F

w

ith

tw

o distinct dorsal sulci)

slightly thicker than previous legs, F

with 1–2 indistinct dorsal sulci, T

i

with one dorsolateral or dorsal

sulcus

thickened, as usual T

i and F

or only one

of them

w

ith 1–2 sulci

Accessory sp

in

e on

leg

15

absent

absent (present only in L

. paghm

anensis)

absent

absent (present only in H

. pervagatus)

Nu

mb

er of sp

urs o

n

fem

ale gon

op

od

s

2+

2 or 3+

3

2+

2

2+

2

fro

m 2 to 5

Dorsod

istal

mod

ification

on

secon

d

fem

ale gon

op

od

al

article

sm

all distodorsal outgrow

th pointed inside

absent (only in L

. giganteus w

ith sm

all rounded

bulge)

probably sm

all distodorsal

outgrow

th pointed inside (L

oksa,

1978: abb. 17)

variable: fro

m sm

all to long styliform

protuberance

Ap

ical claw

of fem

ale

gon

op

od

s (an

d lateral

den

ticles)

sim

ple

variable: usually sim

ple, som

etim

es w

ith lateral

basal denticle

sim

ple

variable: usually sim

ple, som

etim

es

with sm

all or stout accessory denticle

on outer face of claw

Sou

rces

Attem

s 1926, V

erhoeff 1934b, personal

data

Zalesskaja 1978, E

ason 1986a, b, personal data

Verhoeff 1930, L

oksa 1978,

personal data

Zalesskaja 1978, personal data

* T

he giganteus-group of L

ithobiu

s subgenus E

zem

bius C

ham

berlin, 1919 allocated by E

.H

. E

ason for four C

entral A

siatic species: L

. giganteus, L

. pappi E

ason, 1986, L

. haarlovi E

ason, 1986 and L

.

pa

ghm

an

en

sis E

ason, 1986. H

ow

ever, in our opinion, L

. giganteus is very different fro

m other m

em

bers of the group prim

arily by the presence of the secondary sexual m

odification of the tergites in m

ales.

** S

om

e species of the genus H

essebius, w

ith C

entral A

sian distribution, have functionally biarticulated tarsi of legs and big body sizes: H

. perelae Z

alesskaja, 1978, H

. plum

atus Z

alesskaja, 1978, H

.

procurvus Z

alesskaja, 1978 and H

. pervagatus Z

alesskaja, 1978.


Disphaerobius svenhedini (Verhoeff, 1934) comb. nov.

Figs 1–21, 29

Pterygotergum svenhedini—Verhoeff, 1934b: 29, Taf. 4, Abb. 1a, 2a, 3a, 4a, 5a, 6a, 7a (♂, ♀).

Material examined: Type material: Lectotype male (ZSM, in four slides labelled as No. A20030926. Lectotype

1. Pterygotergum svenhedini Verh. ♂ Kopf. Thian-Schan; No. A20030927. Lectotype 2. Pterygotergum svenhedini

Verh. ♂ 1, 2 B. Thian-Schan; No. A20030928. Lectotype 3. Pterygotergum svenhedini Verh. ♂ 3.–8. B. Thian-

Schan; No. A20030929. Lectotype 4. Pterygotergum svenhedini Verh. ♂ 9.–15. B. Thian-Schan).

This male was labeled as lectotype, but since this designation seems to have never been published, it becomes

validated only with the present paper.

Other material examined: Kazakhstan: 1 male, 1 female (ZMUM), 2 females (PSU, No. 555), Almaty

Region, Zhambyl District, 15 km NNW of Karabastau village, Tyrnakty, N 43˚53′, E 75˚30′, stony mountain steppe

with rocks, 850–950 m a.s.l., 20–22.IV.2016, leg. A.A. Fomichev. Mongolia: 1 male (PSU, No. 554), Baitag Bogd

Uul Mt. Range, Gushoot-Shineetijn-Gol river terrace, N 45˚16′, E 91˚04′, stony terrain with sparse Achnatherum

splendens growth, 1900 m a.s.l., 23–26.V.2015, leg. A.A. Fomichev.

Diagnosis. Male D. svenhedini shows the most strongly broadened T 12 divided into two lobes and serrate at

the lateral margin (Figs 6, 16, 18 & 29). In the female, T 12 is noticeably, but not so much broadened as in the male,

strongly notched at the rear edge to form lobes, and the lateral margin is devoid of serration (Figs 4 & 21).

Description. Material examined: Male. Body 28.0 (Mongolia) to 35.5 mm long (Kazakhstan), poorly and

sparsely punctate.

Coloration in alcohol yellow-brownish, with a vague, darker, broad, axial stripe on tergites. Ocellar area dark

(Fig. 7).

Forcipular tergite slightly narrower than cephalic plate, with a ratio of 7.8:7.3. TT 10–Tim broadened, serrate

and infuscate at lateral or all margins (Figs 16 &18). T 12 very strongly broadened so that its lateral parts forming

rounded wing-like outgrowths, strongly serrate at edges. Serration irregular, strongly expressed on sides, gradually

decreasing towards rear edge, fully lacking towards the middle of posterior edge of T 12. Short setae visible at high

magnification between serrations (Fig. 6).

Axial length of T 12, 3.1 mm in both specimens, width 7.6 mm or 8.7 mm. Tergites 10–Tim broadened, serrate

and infuscate at all margins (Figs 16 &18). Tergite 12 especially strongly broadened so that its lateral parts forming

rounded wing-shaped projections distinctly serrate at edges; serration irregular, better developed laterally than

caudally, totally disappearing towards the middle of caudal edge; short setae visible between teeth at high

magnification (Fig. 6). Hind margin of Tim strongly emarginated in front considerably covered by a broadened T

14.

Punctation of cephalic plate denser than of tergites. Cephalic plate slightly broadened, 3.85 mm long, 3.90 mm

wide in both specimens (Figs 1 & 19).

Antennae composed of 18+9 and 20+17 segments, respectively (damaged in both specimens; according to the

original description, antennae 20-segmented, but the lectotype has broken antennae), short, barely reaching the

middle of T 5. Antennomeres elongated, first three covered with sparse and very short setae. Starting with basal 4–

5 antennomeres, antennae bright brown (Fig. 19), densely covered with very small and erect setae.

Ocelli 5–7, posteriormost ocellus always the largest.

Tömösváry’s organ very small, less than half the size of adjoining ocelli (Fig. 7).

Forcipules dark brown, especially at joints. Lateral edges of trochanteroprefemur and part of coxosternite

extended beyond cephalic plate, inner part of trochanteroprefemur concave, with a ventrally bent chitinous rib (Fig.

17). Forcipular coxosternite broad, with 2+2 obtuse and short teeth, shoulders of coxosternite varying from almost

straight to slightly sloped, porodonts stout and strong, about as long as teeth; median notch from weak to wanting

(Figs 9 & 12).

Legs: 14–15P slightly incrassate. 15F incrassate, with two distinct sulci, i.e. a poorly expressed dorsolateral

sulcus and a deep dorsal sulcus, the latter forming apically a round tubercle supporting a cluster of thick and short

setae (Fig. 5). Length of legs: pair 1—t+P 1.00–1.05, F 0.90–0.95, Ti 1.05–1.15, Ts1 0.85–0.95, Ts2 0.45–0.50;

pair 14—t+P 1.60–2.05, F 1.85–2.05, Ti 2.10–2.30, Ts1 1.80–1.85, Ts2 0.80–0.85; pair 15—t+P 1.85–2.20, F 2.25–

2.30, Ti 2.40–2.60, Ts1 2.05–2.20, Ts2 0.80–1.05. Leg plectrotaxy as in Table 2.


FIGURES 1–5. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (1–4 from Kazakhstan, 5 from Mongolia). 1, male front body part (dorsal view); 2, female coxal pores of legs 12–15; 3, male gonopod (ventral view); 4, female TT 10–12; 5, male 15F (lateral view). Scale: 1 mm.


FIGURES 6–9. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (all from Kazakhstan). 6, male T 12, right blade (dorsal view); 7, male ocelli and double circled Tömösváry’s organ (lateral view); 8, male S 15 and 15C; 9, male forcipular coxosternite (ventral view). Scale: 1 mm.


FIGURES 10–15. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (all from Kazakhstan).10, female gonopod (ventrolateral view); 11, female gonopod (dorsolateral view); 12, female forcipular coxosternite (ventral view); 13, female gonopod (lateral view); 14, female gonopod segment I (from inside); 15, female genital sternite and gonopods. Scale: 10–14—0.5 mm; 15—1 mm.


FIGURES 16–20. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (16, 20 from Kazakhstan, 17–19 from Mongolia). 16, male rear body part (dorsal view); 17, male head (ventral view); 18, male rear body fragment (dorsal view); 19, male front body fragment (dorsal view); 20, male rear body fragment (ventral view). Without scale.


TABLE 2. Plectrotaxy of Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. Dashes indicate absence of spines,

brackets—spines on one leg of pair or in one specimen, subscripts –accessory spines.

All sternites sparsely punctate. SS 13–15 and coxae of the same legs covered with very small and dense setae

in rear parts, especially so coxae 15. Attachment points of setae pigmented, therefore fields of setae resembling

brown spots.

Genital sternite short and broad, covered with longer setae. Gonopod 1-segmented, with numerous setae (about

14–17) (Fig. 3).

Coxal pores present on legs 12–15, small, rounded, separated from one another by distances 2–2.5 times

greater than their own diameter; formula 5,4,4,4.

Female. Body 37, 39 and 41 mm long. Diagnostic characters and plectrotaxy as in male, but all tergites and

legs unmodified.

Head broadened, its length and width 3.60 (3.40–3.90) mm and 3.78 (3.50–4.25) mm, respectively.

Width of forcipular tergite 3.85 mm.

T 12 non-serrate, but as in male broadened and lobe-shaped at rear edge, lateral sides subparallel (Figs 4, 21);

width 4.4 (3.90–5.15) mm, length in the middle 2.78 (2.60–3.10) mm. Hind margin of Tim strongly notched (Fig.

21).

Genital sternite broadened, sometimes pigmented (Fig. 15).

Antennae with 20+20 segments, not reaching the middle of T 5.

Ocelli 5–6 on each side of head, varying in size as in Fig. 7.

Forcipules as in male, coxosternite with 2+2 teeth (in one female 3+3) (Fig. 12). Inner face of forcipular

trochanteroprefemur less strongly concave than in male and slightly shorter in length.

Length of legs: pair 1, t+P 1.13 (1.00–1.25), F 0.90 (0.80–1.00), Ti 1.05 (0.95–1.15), Ts1 0.85, Ts2 0.50 (0.45–

0.55); pair 14, t+P 2.01 (1.80–2.20), F 1.75 (1.55–1.90), Ti 1.97 (1.85–2.05), Ts1 1.80 (1.70–1.85), Ts2 0.83 (0.80–

0.85); pair 15, t+P 2.25 (2.05–2.45), F 2.21 (2.10–2.35), Ti 2.46 (2.40–2.55), Ts1 2.17 (2.15–2.20), Ts2 1.05

(1.05–1.10). Plectrotaxy as in Table 2.

Coxal pores as in Fig. 2.

Gonopods with thin setae and 3+3 (Figs 10, 13 & 14), 2+3 and 3+4 unequal spurs (usually obviously 3+3).

First segment with numerous setae on external and, slightly, internal surfaces (Figs 13, 14). Second segment with a

small distodorsal outgrowth shifted mesad (Figs 10, 11). No dorsolateral setae, but 1–2 short setae on first segment

(absent from one specimen), two irregular rows of thin setae on second segment, and a group of very thin and dense

setae on third segment (Figs 11, 13). Gonopodal claw simple, but in two females with a very small, additional

denticle observed on external face of claw closer to its base, probably due to just a slightly uneven outer margin of

claw (Fig. 13).

Remarks. The males at our disposal are differing from the original description (Fig. 29), as well as from the

lectotype (despite the latter’s deformation during mounting on microscopic slides) by showing less sharp rear

edges of T 12 (Figs 6, 16 & 18) and a different plectrotaxy pattern (pair 1: 221/333; pair 2: 222/333; pair 12: 00222/

00333; pair 15: 10310/01432). The presence of four ventral spines on 15P in the original description is given

without disposition details, which may mean either the presence of additional spines or as a misinterpretation.

Lectotype spinulation is much more similar to our specimens (pair 1: 322/233, pair 15: 10311/01333), although

Leg pairs ventral dorsal

t P F Ti C P F Ti

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2–8 – mp amp amp – amp a p a p

9–10 – mp amp amp (a) amp a p a p

11 – (a)mp amp amp (a) amp a p a p

12 – amp amp amp a amp (a) p a p

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14 m amp amp amp a amp p (a) (p)

15 m a(2)

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spines are missing and often visible only as insertion points. In addition, both of the new specimens differ each

other in the shape of TT 12 and 14 (Figs 16 & 18). The deep sulcus on 15F in the individual from Mongolia is

brown at the bottom. However, all these minor variations in secondary sexual characters seem to be intraspecific,

rather depending on age and condition (Fusco et al. 2015).

The female described by Verhoeff (1934b) differs significantly from our specimens, but we have failed to

access the female paralectotypes. First of all, the female he mentioned showed the gonopods with 2+2 spurs: “Die

Gonopoden des ♀ sind denen der Abb. 11 ziemlich ähnlich, also die zwei Genitalklauen kegelig und schräg gegen

einander gestellt, etwa um die eigene Breite von einander getrennt. Genitalklaue am Ende mit einer scharfen

Spitze, aber schräg abgeschnitten, wodurch ein stumpfer Winkel und damit die Andeutung einer 2. Spitze

entsteht…” [Gonopods of the ♀ are very similar to those depicted in fig. 11, so that the two gonopodal spurs are

tapered and obliquely juxtaposed, separated by their own width. Gonopodal claw at the end with a sharp point, but

cut obliquely, creating an obtuse angle and thus the forming as if a 2nd tip…] (Verhoeff 1934b: 31). Despite this,

Verhoeff did not mention a distodorsal outgrowth on the second gonopodal segment. Moreover, Verhoeff, when

describing the female gonopod, referred to a figure of a different species, i.e. Lithobius giganteus (= Lithobius

mongolicus Verhoeff, 1934) (Verhoeff 1934b: Taf. 5, Abb. 11a).

Distribution (Fig. 30). China: Xinjiang Uygur Autonomous Region, Urumqi (Tian Shan) (Verhoeff 1934b).

Kazakhstan: Almaty Region, Zhambyl District. Mongolia: Baitag Bogd Uul Mt. Range.

Disphaerobius loricatus (Sseliwanoff, 1881) comb. nov.

Figs 22–26

Lithobius loricatus—Sseliwanoff, 1881: 16 (♂).Lithobius magnificus—Trotzina, 1895: 108, Taf. I: figs 1–4 (♂) syn. nov.Disphaerobius magnificus—Attems 1927: 243 (new comb.).Lithobius loricatus—Zalesskaja 1978: 119 (♂); Farzalieva & Zalesskaja 2003: 265, 266: figs 1–13 (♂♂, ♀♀).

Material examined: Type material: Holotype male (ZIL, No. 94), labelled in Latin “Lithobius loricatus Ssel. ♂,

inter Semipalatinsk et Ajagus, leg. A. Schrenk” = Kazakhstan, East Kazakhstan Region, between Semey and

Ayagoz, no date, leg. A. Schrenk. Allotype: 1 female (ZMUM), Russia, Orenburg Area, Sol-Iletsk District,

Chybynda, base of limestone denudation, slope, VI.2000, leg. S.L. Esyunin & G.Sh. Farzalieva. 2 males, 1 female

(PSU, No. 116), same locality, limestone plateau, saline land with Astragalus, VI.2000, leg. S.L. Esyunin & G.Sh.

Farzalieva.

Other material examined: Russia: Orenburg Area: 1 male, 2 females (ZMUM), Sol-Iletsk District,

Chybynda, base of limestone denudation, slope, limestone plateau, salina with Astragalus growth, VI.2000, leg.

S.L. Esyunin & G.Sh. Farzalieva; 1 male, 1 female (PSU, No. 179), 1 male (PSU, No. 147), same locality,

limestone slope, V–VI.2003, leg. T.K. Tuneva; 1 male, 3 females (ZMUM), same locality, V.2015, leg. S.L.

Esyunin; 1 male, 1 female (PSU, No. 117), near Aituar Village, stony scree under rock, V.1996, leg. T.I. Gridina &

same locality, scree on bank of Ural River, V.2015, leg. S. Dedyukhin; 1 male, 2 females (PSU, No. 450), near

Boevaya Gora Village, steppe, IV.2009, leg. V.O. Kozminykh & same locality, stony steppe, V.2015, leg. S.L.

Esyunin; 1 female (PSU, No. 191), Donguz steppe, under stones, IV.2000, leg. S.L. Esyunin & G.Sh. Farzalieva; 9

males, 6 females (PSU, No. 316), 3 km NW Pervomaiskii Village, steppe, Stipa, VI.2007 & same locality, Donguz

steppe, X.2007–V.2008, V.2009, all leg. V.O. Kozminykh; 1 male, 2 females (PSU, No. 481), Svetlinskii District,

near Dombarovkii Village, steppe, V.2015, leg. S.L. Esyunin; 6 males, 5 females (PSU, No. 448), Sol-Iletsk

District, 8 km SW Troitsk Village, Tasbulak dried-up river-bed, cretaceous sediments, 8–22.VI.2008, leg. V.O.

Kozminykh & V.A. Nemkov. Kazakhstan: 1 female (PSU, No. 483), Aktobe Region, Mugalzhar District, 30 km E

of Emba Village, Mugodzhar Hills, N 48˚46′, E 58˚32′, 550 m a.s.l., no date, leg. A.V. Ivanov.

Diagnosis. In D. loricatus males, T 14 is the broadest, but without two lobes (Figs 22 & 26). In females, T 12

is of similar length and width, concave at the rear edge, without formed lobes (Farzalieva & Zalesskaja 2003: Fig.

10). The main differences between D. svenhedini and D. loricatus are summarized in Table 3.

Description. See a detailed redescription by Farzalieva & Zalesskaja (2003).


FIGURES 21–24. Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (21) from Kazakhstan and D. loricatus(Sseliwanoff, 1881) comb. nov. (22–24) from Chybynda, Orenburg Area, Russia. 21, female rear body fragment (dorsal view); 22, male rear body fragment (dorsal view); 23, male front body fragment (dorsal view); 24, male head (ventral view). Without scale.


FIGURES 25–29. Disphaerobius loricatus (Sseliwanoff, 1881) comb. nov. (25–28) from Chybynda, Orenburg Area, Russia and D. svenhedini (Verhoeff, 1934) comb. nov. (29) from China, type locality. 25, male 15F (dorsolateral view); 26, male TT 13–14 (dorsal view); 27, male ocelli and Tömösváry’s organ (indicated by arrow) (lateral view); 28, female gonopod (dorsolateral view); 29, male TT 11–14 (dorsal view) (without scale, after Verhoeff (1934b)). Scale: 25–26—1 mm; 27–28—0.5 mm.

Remarks. The head and tergites are strongly punctate; the forcipules broadened, significantly extended

beyond the cephalic plate (Fig. 23), the trochanteroprefemur is slightly concave on the inner surface, with an

almost straight, ventral, chitinous rib (Fig. 24). All males have T 14 and Tim broadened and serrate, especially

strongly broadened is T 14, their surface roughly rugose (Figs 22 & 26). In addition, some specimens show serrate


edges of T 13 and slightly serrate posterolateral edges of T 12. In all males, 15F are thickened, with two distinct

sulci, i.e. a poorly expressed dorsolateral sulcus and a deep dorsal sulcus, the latter forming apically a round

tubercle supporting a cluster of thick and short setae; some specimens in addition with 1 or 2 poorly expressed

dorsal sulci on 15P; 15Ti with a barely visible dorsolateral sulcus. In most males, very fine and dense setae cover

rear halves of SS 14–15, as well as 14C, 13P, F and Ti ventrally. Tarsi 1–13 with two ventral tarsal pectens.

TABLE 3. Main morphological differences between Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. and D.

loricatus (Sseliwanoff, 1881) comb. nov.

Distribution (Fig. 30): Steppe zone from the Sol-Iletsk District, Orenburg Area, Russia in the west to the East

Kazakhstan Region of Kazakhstan (Sseliwanoff 1881) in the east.

FIGURE 30. Distribution of Disphaerobius svenhedini (Verhoeff, 1934) comb. nov. (circle) and D. loricatus (Sseliwanoff, 1881) comb. nov. (square). Red coloration indicates type localities.

Discussion

The sex of lithobiomorph centipedes is always easy to recognize, based on morphological traits, as primary sexual

dimorphism is related to the shape of the gonopods. Secondary sexual dimorphism is well-visible mainly through

the presence of sulci, hollows, pits, grooves, processes, tufts of setae, swellings and/or other modifications on male

legs 15, sometimes 14, rarely also 13 (Eason 1993). Nevertheless, there are also some exceptions, in which

secondary sexual dimorphism is not restricted to legs (Crabill 1960).

Disphaerobius svenhedini (Verhoeff, 1934) comb. nov.

Disphaerobius loricatus (Sseliwanoff, 1881) comb. nov.

Male forcipules trochanteroprefemur with a big notch on inner edge (Fig. 17)

trochanteroprefemur with a small notch on inner edge (Figs 23, 24)

Macrotergites macrotergites starting from 10 strongly carved on hind margins, forming blades

macrotergites only with small cuttings on hind margins, without blades

Male modified tergites

TT 10–Tim; most modified T 12: with big saw-like serrate wing-like outgrowths, forming clear margin between blades in the middle. Tergite surface almost smooth, only slightly punctated (Figs 6, 16, 18 & 29)

TT 14 and Tim (sometimes 12 and 13); most modified T 14: thickened irregularly serrate lateral edges, without blades on the rear edges. Tergite surface strongly wrinkled (Figs 22, 26)

Female gonopods with 3+3 (2+2 in original description) with 2+2 spurs


Examples of sexual dimorphism related to the anterior part of the body are represented by the European

Lithobius muticus C.L. Koch, 1847, in which the males have a relatively much wider head than the first tergite in

comparison with the females (Koren 1992), and perhaps Eupolybothrus dolops Zapparoli, 1998, with broadened T

1 and partly modified forcipulae in the males (but no females are known yet). A far more bizarre example concerns

the North American Paitobius zinus (Chamberlin, 1911): males have elongated and distorted forcipulae compared

to the females (Crabill 1960). Males of D. svenhedini also show larger forcipulae with a longer and more strongly

curved trochanteroprefemur than females do, but the expression of such sexual dimorphism is rather weak and is

evident only in fully grown individuals. Other cases of sexual dimorphism are related to the posterior part of the

body. In males of Pleurolithobius Verhoeff, 1899, the intermediate tergite is endowed with prolonged posterior

projections (Berlese 1894; Zapparoli 1989; Zapparoli & Minelli 1993). The posterior sternites and the coxae of the

last legs of male Pleurolithobius species are covered with short and dense setae. Male T 14 of Lithobius

(Dacolithobius) domogledicus Matic, 1961 is elongate posteriorly and covered with long strong setae, in contrast to

an unmodified tergite in the females (Matic 1966).

In several species of the genus Gosibius Chamberlin, 1912, males have serrate lateral sides of TT 7–10, which

are broadened as well (e.g. Gosibius fusatus Chamberlin, 1941 and probably also G. intermedius Chamberlin, 1917,

of which females are not known yet). In the species Atethobius mirabilis Chamberlin, 1915 and Atethobius scutiger

Verhoeff, 1934 (Verhoeff 1934a), both of which are known only from males, T 14 is greatly enlarged and

subcircular. Nevertheless, sexual dimorphism in Disphaerobius as observed in serrate and broadened male TT (10–

)13–Tim seems be to the most ostentatious among lithobiomorph centipedes.

Secondary sexual dimorphism is usually related to sexual selection, when more picturesque males have a

higher probability to charm females and increase fitness (e.g. peacocks). Although these peculiar structures can be

suggested as being related to mating (Lewis 1981), neither experimental evidence nor direct observations of mating

behaviour have been published yet as regards lithobiomorphs.

Acknowledgements

The authors wish to thank Dr. S.I. Golovatch (Moscow, Russia) for editing an advanced draft. The first author is

most grateful to Dr. S.L. Esyunin (Perm, Russia) for his constant guidance, encouragement and support. The

authors are grateful to Dr. A.V. Grischenko, Head of the Zoological Museum of the Perm State University, for

assistance in use the SEM facilities. We are very much obliged to Dr. G. Edgecombe (Natural History Museum,

London, UK) who kindly edited an advanced draft for English usage. Our deepest gratitude extends to A.A.

Fomichev, A.V. Ivanov and V.O. Kozminykh who donated us their material for the present study. Special thanks are

also addressed to T.K. Tuneva and Yu.V. Dyachkov for provided material, as well as to S. Friedrich (ZSM, Munich,

Germany) for the loan of the type material of D. svenhedini. We are grateful to Dr. L. Dányi (Hungarian Natural

History Museum, Budapest, Hungary) who created the distribution map.

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AbstractIntroductionMaterial and methodsTaxonomic partDisphaerobius Attems, 1926Disphaerobius svenhedini (Verhoeff, 1934) comb. nov.Disphaerobius loricatus (Sseliwanoff, 1881) comb. nov.DiscussionAcknowledgementsReferences

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