European Journal of Taxonomy 227: 1-19

http://dx.doi.org/10.5852/ejt.2016.227

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2016 • Vagalinski B. & Golovatch S.I.

Research article

urn:lsid:zoobank.org:pub:41346283-D50E-4A52-A6AE-2FA5ED23FlF4

Two new species of Anamastigona from Cyprus and an updated

key to species of the genus (Diplopoda: Chordeumatida:

Anthroleucosomatidae)

BOYAN VAGALINSKI 1 * & SERGEI I. GOLOVATCH 2

1 Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences,

2 Yurii Gagarin Street, 1113, Sofia, Bulgaria,

institute for Problems of Ecology & Evolution, Russian Academy of Sciences,

Leninsky prospect 33, Moscow 117051, Russia.

Email: [email protected]

2 Email: [email protected]

1 urn:lsid:zoobank.org:author:973B8725-039E-4F29-8D73-96A7F52CF934

2 urn:lsid:zoobank.org:author:71532F45-BDD5-415D-BC54-86256E5D5D4A

Abstract. Two new species of the millipede genus Anamastigona are described, A. cypria sp. nov. and

A. strasseri sp. nov., both from the island of Cyprus. The new species are considered to be most similar

to A. terraesanctae Golovatch & Makarov, 2011 from Israel, the three of them forming a species group

which is briefly characterized. An updated key to all 20 recognized species of the genus Anamastigona

is provided and a short overview of the taxonomy of the genus is given.

Keywords. Millipedes, taxonomy, distribution, endemics, synonyms.

Vagalinski B. & Golovatch S.I. 2016. Two new species of Anamastigona from Cyprus and an updated key to

species of the genus (Diplopoda: Chordeumatida: Anthroleucosomatidae). European Journal of Taxonomy 227:

1-19. http://dx.doi.org/10.5852/eit.2016.227

Introduction

The millipede germs Anamastigona Silvestri, 1898 (Chordeumatida: Anthroleucosomatidae) is hitherto

known to comprise 17-19 species native to the Apennine and Balkan peninsulas, several Mediterranean

islands, the northwestern Aegean coast of Turkey, Israel and the Caucasus, with one species, A. pulchella

(Silvestri, 1894), showing a broader, scattered distribution across western Europe, probably a result of

human activity.

The name Anamastigona has a long history dating back to Silvestri (1898), followed by numerous, mostly

single-species descriptions and redescriptions by various authors, until the last two members of the

genus —A. mauriesi and A. terraesanctae - were described by Golovatch & Makarov (2011). The latest

comprehensive work on the genus in the framework of a review of the family Anthroleucosomatidae is

that by Curcic et al. (2008). These authors presented a key to the species of Anamastigona , as part of

a key to the genera and species of Anthroleucosomatidae, and suspected that A. aspromontis (Strasser,

1

European Journal of Taxonomy 227: 1-19 (2016)

1970) and A. halophila Verhoeff, 1940 were junior synonyms of A. meridionalis Silvestri, 1898 and

A. bilselii (Verhoeff, 1940), respectively; both of those synonymies were formally proposed later by

Golovatch & Makarov (2011). However, comparison of the original drawings of A. meridionalis and A.

aspromontis reveals considerable differences between the two species, making their synonymy rather

unlikely (see also Discussion). The synonymy of A. halophila with A. bilselii seems quite probable

considering the known distribution of A. bilselii , but nevertheless a direct comparison of type material

is needed to secure the synonymy in question.

The present study adds another two species to Anamastigona , currently representing the only known

members of the genus and of the order Chordeumatida in general from Cyprus (Vagalinski et al. 2014).

The existence of the two new species fills in the geographical gap between the Aegean islands and Israel

within the distribution range of Anamastigona. Considering the increased number of species recognized

in Anamastigona , an updated key is provided, in which a total of 20 species is recognized.

Material and methods

All specimens are preserved in 70% ethanol. Gonopods, legs, antennae and vulvae of some specimens

were dissected and mounted on permanent euparal or eukitt slides and observed under an Olympus

BX51 light microscope. Colour micrographs were taken with the aid of a Carl Zeiss Discovery V8 stereo

microscope, with a Nikon Coolpix S3 700 camera mounted on one of the eyepieces. SEM pictures were

obtained with a JEOL JSM-5510 scanning electron microscope after sputter-coating of the prepared

body parts with gold-palladium.

The descriptions of the new species mostly follow the pattern of Golovatch & Makarov (2011).

_ r

Terminology of the gonopod parts and the metrical indices are after Curcic et al. (2008) and Golovatch

& Makarov (2011), with minor changes.

Abbreviations used:

CIX = macrochaetal index

MA = macrochaetal angle

MIX = median index

NHMC = Natural History Museum of Crete, University of Crete, Irakleio, Crete, Greece

NMNHS = National Museum of Natural History, Sofia, Bulgaria

ZMUM = Zoological Museum, State University of Moscow, Russia

Results

Class Diplopoda Blainville-Gervais, 1844

Order Chordeumatida Koch, 1847

Family Anthroleucosomatidae Verhoeff, 1899

Genus Anamastigona Silvestri, 1898

Anamastigona Silvestri, 1898: 661. Type species: Craspedosoma pulchellum Silvestri 1894: 199, by

original designation.

Prodicus Attems, 1899: 317.

Hellasdicus Verhoeff, 1940a: 273.

Paraprodicus Verhoeff, 1940b: 24.

Balkandiciis Strasser, 1960: 112.

Osmandicus Strasser, 1960: 105.

Antrodicus Gulicka, 1967: 2.

2

VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

Diagnosis

A genus of the family Anthroleucosomatidae characterized by a body consisting of 29 segments plus

the telson and by the following gonopodal characters: caudal face of anterior gonopods with lamellar

central part, but without well-developed syncoxite, and with colpoxites; posterior gonopods with well-

developed, strongly chitinized angiocoxites and with usually more or less membranous colpoxites.

Anamastigona cypria sp. nov.

um:lsid:zoobank.org:act:FD1105FF-4E6D-4851-A5B6-477FFDD8DFD2

Figs 1-3

Anamastigona sp. 1 - Vagalinski et al. 2014: 540, fig. 12.

Diagnosis

Resembles its most similar congeners - A. terraesanctae and A. strasseri sp. nov. - by the anterior

gonopods possessing well-developed colpocoxites and an elaborate median lamella, which is more

or less densely covered with long, setiform filaments; and by the posterior gonopods consisting of

large, relatively stout angiocoxites, bearing several setae and rather prominent telopodital rudiments,

and of fine, membranous colpocoxites. Differs from them by being on average slightly smaller, with a

somewhat darker colouration, and by certain details of gonopod structure, namely, the lateral arms of

the anterior gonopods branching distally into 2 processes and by the presence of a subapical unciform

process on the angiocoxites of the posterior gonopods; further morphological differences between the 3

species are summarized in Table 1.

Etymology

The epithet of this new species emphasizes its provenance from Cyprus.

Type material

Holotype

CYPRUS: S (intact), FC0022, Nicosia, Pediaios, stream under Eucalyptus , S4, 27 Nov. 2011-15 Jan.

2012, pitfall traps, Simaiakis, Stylianou & Tambouri leg. (NHMC).

Paratypes

CYPRUS: 3 SS (intact, 1 half-broken in the caudal part), 1 $ (with heavily damaged and fragmented

head and front body, right vulva mounted on slide), same sample as holotype (NHMC); 1 S (in 2

pieces, with broken distal parts of antennae, gonopods mounted for SEM), FC0060, Nicosia, Troodos

Mts, 4 km before Prodromos square, pine forest next to the road, 1305 m, 11 Feb.-19 Mar. 2012, pitfall

traps, Simaiakis leg. (NHMC); 1 S (in 2 pieces, gonopods dissected), FC0019, Nicosia, Athalassas Park

South, corridors with Eucalyptus, SI, 27 Nov. 2011-15 Jan. 2012, pitfall traps, Simaiakis, Stylianou &

Tambouri leg. (ZMUM); 2 SS (one dissected, in 2 parts + pleurotergum 7, with gonopods mounted for

SEM, the other into head, collum and 3 body pieces; gonopods, antenna, legs 1, 3, 5, 7 and 15 mounted

on slide), FC0009, Troodos Mts, 2.5 km after Prodromos square, mixed pine forest, 1615 m, 18 Oct.-26

Nov. 2011, pitfall traps, Simaiakis & Louca leg. (NMNHS).

Description

Measurements. With 30 segments including telson, with 1-2 legless segments; males 7.2-9.2 mm in

length, 0.6-0.65 in height at mid-body and 0.75-0.8 mm ma xim al width; holotype 9.1 mm long; the

female somewhat longer than the males (exact length impossible to measure due to the damaged frontal

part), 0.75 mm high at mid-body, 0.95 mm maximal width (at segments 25-26).

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European Journal of Taxonomy 227: 1-19 (2016)

Colouration. Visibly faded on most specimens. Mostly brown, with scattered, small, bright spots and

veins, giving a marble-like pattern to the tegument; prozonae on posterior segments dorsally blackish;

with smaller, oval, bright spots just above shoulders; sterna yellowish-beige; legs beige to light brown.

External structures. Tegument smooth, only hind margins of pleuroterga very finely and sparsely

striated dorsally. Head with 20-24 ocelli, arranged in 8 oblique rows: 1 +1 + 2 + 3 + 4+4 (5) + 3 (4,5)+2 (3),

counting from hind lower comer of ocellarium. Head moderately setose: vertex, frons, labmm and

mandibular cardines more or less evenly covered with short, stout setae, these latter being somewhat

longer on labmm and distal part of frons. Frons in males flat to gently convex. Antennae (Fig. 2A)

almost 2 times as long as head; antennomeres 2 and 4 subequal, somewhat longer than 6, antennomere

3 ca 2 times longer than 4, slightly shorter than antennomere 5. Fabmm with 3 small, rounded

marginal teeth. Gnathochilarium with a rather large, broad, subtriangular promentum, about half as

long as stipites. Collum smooth, without median groove. In width, collum < segment 2 < 3 < 4 <

head - 5 - 8-18 (19) <6 — 7 (males), after segment 19 increasingly narrowed towards telson. Axial

Fig. 1 . External habitus of Anamastigona cypria sp. nov., paratype from Nicosia, Pediaios. A. Anterior

part of body, dorsal view. B. Anterior part of body, lateral view. C. Middle part of body, lateral view.

D. Posterior part of body, lateral view. Scale bars = 0.5 mm.

4

VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

suture well-developed, narrow. Stricture between pro- and metazonae quite wide and rather shallow.

Paraterga small, somewhat edgy dorso-lateral swellings, forming a mild ridge between lateral and

median macrochaetae; most prominent on segments 8-19, then gradually diminishing towards telson,

completely disappearing on segments 28 and 29. Metatergal setation typical, macrochaetae 0.12-0.2 mm

long, on same segment more or less equal in length; CIX (15) ~ 0.5-0.6, MIX (15) ~ 1.3, MA (15)

~ 135-145°. Pleurotergum 7 unmodified. Epiproct short, broad, truncated caudally, with a straight to

Fig. 2. Non-gonopodal characters of Anamastigona cypria sp. nov., S3, paratypes. A. Antenna (setae

omitted). B. Leg 1. C. Leg 3. D. Leg 5. E. Leg 7 (claw broken off). F. Leg 10. G. Leg 11. H. Leg 24.

Scale bar = 0.2 mm.

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European Journal of Taxonomy 227 : 1-19 ( 2016 )

Fig. 3. Genitalia of Anamastigona cypria sp. nov., paratypes (c?: A—F; ?: G) from Nicosia, Troodos

Mts, 4 km before Prodromos square. A. Posterior gonopods, caudal view. B. Left angiocoxite of the

posterior gonopods, caudal view. C. Same aspect, mid-part strongly magnified. D. Anterior gonopods,

lateral view. E. Anterior gonopods, caudal, slightly lateral view. F. Same aspect. G. Left vulva, caudal

view (setae omitted; receptaculum seminis seen by transparency, but drawn with solid lines for better

emphasis). Abbreviations: a = angiocoxite; ap = anterior process; c = colpoxites of anterior gonopods;

ct = central tube; / = lamellar median part; la = lateral arm; lo = lamellar outgrowth; k = colpoxites of

posterior gonopods; o = opening; pp = posterior process; pt = posterior tube; s = central depression; te =

telopodital rudiment; up = unciform process. Scale bars (drawings) = 0.1 mm.

6

VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

slightly rounded caudal margin; medially with a pair of two long setae, distally with a row of 7 short

setae. Paraprocts with 3 long marginal setae each. Hypoproct semi-elliptic, with 3 long marginal setae.

Male legs. Pairs 1 (Fig. 2B) and 2 significantly shorter and thinner than following ones, mesally with

fine, short setae, forming tarsal brushes; coxa 2 ventrally with a gland opening; pairs 3 (Fig. 2C) and

4 conspicuously thick, each with a stout, shortly bifurcated bump dorso-parabasally on prefemur; 5-7

increasingly longer, 5 (Fig. 2D) with a short, thorn-like process anteriorly on prefemur, 7 (Fig. 2E) with

an extremely long tarsus, the latter more than 2 times as long as postfemur + tibia; coxa 10 (Fig. 2F)

ventrally with small tubercle protruding from gland opening; coxa 11 (Fig. 2G) ventrally with blunt

process; following pairs without modifications; leg 24 (Fig. 2H) 1.25-1.4 times as long as mid-body

height, its tarsus 1.8-1.9 times longer than postfemur + tibia.

Gonopods (Fig. 3A-F). Anterior gonopods (Fig. 3D-F) fused in their proximal parts, consisting of two

lateral arms (la), each divided distally into two processes - a larger posterior (pp) and a smaller anterior

one (ap), both processes tapering and bent caudad; a lamellar median part (l), densely covered with long

setiform filaments; and two elongated colpocoxites ( c ), each ending apically as lamellar outgrowths (lo),

basally surrounding a subtriangular central depression (s). Posterior gonopods (Fig. 3A-B) reduced;

angiocoxites (a) elongated, relatively stout, somewhat tapering, subapically bearing thin, unciform

process (up) bent caudad, caudal face microsquamate, bearing several setae; telopodital rudiments (te)

lobe-like, pigmented, positioned caudo-laterobasally on a, colpoxites (k) placed close to each other,

considerably shorter than a, slender, weakly chitinized.

Vulva (Fig. 3G). Compact, box-lilce, somewhat meso-laterally compressed; opening (o) positioned

mostly apically on bursa, narrow; operculum equal to bursa, with shallowly bifurcated apex; setation

rather short, present only on distal parts of bursa and operculum. Receptaculum seminis double,

consisting of larger, somewhat bent posterior tube (pi) and smaller, more or less straight central tube (ct).

Anamastigona strasseri sp. nov.

urn:lsid:zoobank.org:act:07A0082F-06B5-4078-96Cl-4B0A9B3DA43B

Figs 4-6

Anamastigona sp. 2 - Vagalinski et al. 2014: 541, fig. 12.

Diagnosis

Resembles its most similar congeners -A. terraesanctae and A. cypria sp. nov. - by the anterior gonopods

possessing well-developed colpocoxites, and an elaborate median lamella, which is more or less densely

covered with long, setiform filaments; and by the posterior gonopods consisting of large, relatively stout

angiocoxites, bearing several setae and rather prominent telopodital rudiments, and of fine, membranous

colpocoxites. Differs from them mostly by the lateral arms of the anterior gonopods being blunt and

barely bent caudad, rather than tapering and strongly bent caudad. Further morphological differences

between the 3 species are summarized in Table 1.

Etymology

This species is named in memory of Karl Strasser (1903-1981), a prominent myriapodologist who,

among other things, described several species of Anamastigona.

Type material

Holotype

CYPRUS: S (in 3 parts: head + first 6 segments, segment 7 and rest of body; half-broken behind

segment 18), FC0003, Paphos, Pano Panagia village, next to a winery, patches with Eucalyptus, 800 m,

hand collection, 8 Nov. 2011, S. Simaiakis leg. (NHMC).

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European Journal of Taxonomy 227 : 1-19 ( 2016 )

Description

Measurements. Body segments 28+1+T, 8.2 mm long, 0.7 mm high at mid-body, 0.85 mm ma xim al

width.

Colouration. Marbled brownish (somewhat more contrasting compared to A. cypria sp. nov.), more

intense on posterior half of body; with blackish transverse stripes dorsally on prozonae; ventral side

and legs beige to light brown; large, light, cloud-like spots present dorso-laterally just above shoulders.

External structures. Tegument smooth, only hind margins of pleuroterga finely striolate dorsally. Head

with 19-20 blackish ocelli arranged in 8 oblique rows: 1 +1+2 + 3 (2)+ 3+4+ 3 (4)+ 2(3), counting

from hind lower comer of ocellarium. Head rather sparsely setose: vertex and upper part of frons bare,

Fig. 4. External habitus of Anamastigona strasseri sp. nov., holotype. A. Head with first 7 segments,

lateral, slightly dorsal view. B. Head with first 7 segments, dorsal, slightly lateral view. C. Segment 8 to

telson, lateral view. Scale bars = 0.5 mm.

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VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

lower part of frons and labrum moderately densely covered with two types of setae - some very short

ones, the others ca 2 times as long; several very short setae on occiput. Frons flat or very gently convex.

Antennae almost 2 times as long as head; antennomeres 2, 4 and 6 subequal, antennomere 3 ca 2 times

longer, slightly shorter than antennomere 5. Labrum tridentate. Gnathochilarium with rather large,

triangular, somewhat broader than long promentum; stipites with several long lateral setae. Collum with

median longitudinal groove. In width, collum < segment 2 < head ~ 3 < 4 < 5 ~ 8-18 (19) < 6 < 7; after

segment 19 increasingly narrowed towards telson. Axial suture well-developed, thin. Stricture between

pro- and metazonae quite broad and rather shallow. Paraterga small, rounded, dorsolateral swellings,

most prominent on segments 8-19, then gradually diminishing towards telson, completely disappearing

on segments 28 and 29. Metatergal setation typical (macrochaetae mostly broken off), macrochaetae

0.2-0.25 mm long. CIX (15) ~ 0.6, MIX (15) ~ 1.6, MA(15) ~ 135°. Pleurotergum 7 almost unmodified,

its ventral margin forming very low ridge. Epiproct truncated caudally, caudal margin with several

small, rounded, scale-like protuberances. Each paraproct with single long marginal seta. Hypoproct

semi-elliptic, with two long marginal setae.

Legs. Pairs 1 and 2 shortest, with very thin prefemur and femur, mesally with tarsal brushes; coxa 2 with

gland opening; pairs 3 and 4 (Fig. 5A) very thick, with stout, shortly bifurcated bump dorso-parabasally

on pre femur; 5-7 increasingly longer, tarsus of 7 (Fig. 5B) very long, ca 1.6 times as long as postfemur +

tibia; coxa 10 (Fig. 5C) ventrally expanding into cylindrical process, expanding apically into somewhat

curved, foam-like protrusion; coxa 11 (Fig. 5D) with two processes: a proximal, ventral one, similar to

that on coxa 10, the other a distal, ventro-anterior, short and blunt, process; leg 24 ca 1.3 times as long

as mid-body height, its tarsus ca 2 times as long as postfemur + tibia.

Gonopods (Fig. 6A-D). Anterior gonopods (Fig. 6A-B) similar to those of A. cypria sp. nov., the main

difference being the lateral arms (la), which are uniramous, apically blunt/narrowly rounded, barely

bent caudad. Posterior gonopods (Fig. 6C-D) reduced; angiocoxites (a) stout, flattened apically,

microsquamate, caudo-laterally with several setae; telopodital rudiments (te) short, rounded, composed

of 2 articles, pigmented, positioned caudo-latero-parabasally on a; colpoxites (k) placed almost one

behind the other rather than side by side, slender, somewhat lower than a.

Fig. 5. Legs of Anamastigona strasseri sp. nov., $, holotype. A. Leg 4. B. Leg 7. C. Leg 10. D. Leg 11.

Scale bar = 0.2 mm.

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European Journal of Taxonomy 227: 1-19 (2016)

Fig. 6. Gonopods of Anamastigona strasseri sp. nov., S, holotype. A. Anterior gonopods, caudal, slightly

lateral view. B. Anterior gonopods, frontal, slightly lateral view. C. Posterior gonopods, frontal view.

D. Right colpoxite of posterior gonopods, caudal view. Abbreviations as in Fig. 3. Scale bar = 0.1 mm.

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VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

Table 1. Diagnostic characters of Anamastigona cypria sp. no v.,A. strasseri sp.

Golovatch & Makarov, 2011.

nov. and A terraesanctae

A. cypria sp. nov.

A. strasseri sp. nov.

A. terraesanctae

Lateral arms of

distally divided into

uniramous

anterior gonopods

2 processes

Angiocoxites of

posterior gonopods

subapically bearing an

unciform process

ending blunt, without

processes

subapically bearing a

finger-like process

Head

broader than segment 2

more narrow than

segment 2

Collum

without groove

with a median groove

without groove

MIX

1.3

1.6

1.3

Marginal setae on

each paraproct

3

1

3

Marginal setae on

hypoproct

3

2

Tarsus 7

ca 2 times as long as

postfemur + tibia

ca 1.6 times as long as

postfemur + tibia

ca 2 times as long as

postfemur + tibia

Coxa 10

with a small tubercle

protruding from a gland

opening

with a process ending in a

foam-like protrusion

with a gland opening;

without processes

Remarks

The position of the posterior gonopod colpocoxites in this species is unusual - they are almost completely

oriented one behind the other, rather than next to each other, i.e., more perpendicular than parallel to the

sternal axis. However, in the presence of only one examined male, this peculiarity may represent nothing

more than individual variation.

Comparative material

Anamastigona terraesanctae Golovatch & Makarov, 2011

Fig. 7

Material examined

ISRAEL: 1 $ (gonopods prepared for SEM), 2 $ $, Shuni Park, 1 km N of Binyamina, 32°32' N, 34°57' E,

60 m a.s.l., date unknown, Yu. Marusilc leg. (ZMUM).

Remarks

Gonopods (Fig. 7A-E). Anterior gonopods (Fig. 7C-E) consisting of two lateral, uniramous arms

(la), ending with strongly bent, unciform tips; a lamellar median part (/), relatively sparsely covered

with setiform filaments; and two robust colpocoxites (c), basally surrounding a subtriangular central

depression (s). Posterior gonopods (Fig. 7A-B) consisting of stout, clavate angiocoxites (a), caudo-

latero-basally bearing short, rounded telopodital rudiments (te); and of slender, weakly chitinized

colpoxites (k).

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European Journal of Taxonomy 227: 1-19 (2016)

Key to known species of Anamastigona based on gonopodal characters

1. Posterior gonopods with telopodital rudiments .2

- Posterior gonopods either without telopodital rudiments or these only represented by pigmented

spots .14

2. Telopodital rudiments of posterior gonopods positioned apically on angiocoxites.

. A. mauriesi Golovatch & Makarov, 2011

- Telopodital rudiments of posterior gonopods positioned basally/parabasally on angiocoxites.3

Fig. 7. Gonopods of Anamastigona terraesanctae Golovatch & Makarov, 2011, c?, non-type from Israel,

the vicinity of Binyamina. A. Posterior gonopods, frontal view. B. Apical part of left angiocoxite, frontal

view. C. Anterior gonopods, caudal, slightly lateral view. D. Anterior gonopods, caudo-lateral view.

E. Anterior gonopods, lateral, slightly caudal view. Abbreviations as in Fig. 3.

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VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

3. Anterior gonopods with flagella. A. matsakisi Mauries & Karamaouna, 1984

- Anterior gonopods without flagella.4

4. Angiocoxites of posterior gonopods with setae.5

- Angiocoxites of posterior gonopods (excluding telopodital rudiments) without setae.9

5. Angiocoxites of posterior gonopods reaching approximately same level as both colpocoxites and

telopodital rudiments. A. meridionalis Silvestri, 1898

- Angiocoxites of posterior gonopods considerably outreaching both colpocoxites and telopodital

rudiments.6

6. Caudal face of anterior gonopods with neither setae nor fringes, nor spiculi.

. A. hispidula (Silvestri, 1895)

- Caudal face of anterior gonopods more or less densely pilose/spiculate.7

7. Lateral arms of anterior gonopods more or less straight, blunt apically. A. strasseri sp. nov.

- Lateral arms of anterior gonopods bent strongly caudad, tapering apically.8

8. Colpocoxites of anterior gonopods short and robust; angiocoxites of posterior gonopods massive,

somewhat bulging apically. A. terraesanctae Golovatch & Makarov, 2011

- Colpocoxites of anterior gonopods elongated, with membranous ends; angiocoxites of posterior

gonopods rather slender, somewhat narrowing distally. A. cypria sp. nov.

9. Angiocoxites of posterior gonopods very slender, finger- or horn-shaped.10

- Angiocoxites of posterior gonopods stouter, more massive.12

10. Apical parts of lateral arms of anterior gonopods directed mesad.

. A. radmani Makarov et al. , 2007

- Apical parts of lateral arms of anterior gonopods directed laterad.11

11. Colpocoxites of posterior gonopods mostly to entirely fused into a single plate .

. A. pulchella (Silvestri, 1894)

- Colpocoxites of posterior gonopods well separated. A. bilselii (Verhoeff, 1940)

12. Telopodital rudiments of posterior gonopods well-pronounced lobes.

. A. aspromontis (Strasser, 1970)

- Telopodital rudiments of posterior gonopods represented by rounded, wealdy protruding bumps

.13

13. Telopodital rudiments of posterior gonopods with one seta each.

r

. A. mediterranea Curcic, Makarov & Lymberakis, 2001

- Telopodital rudiments of posterior gonopods without setae. A. penicillata (Attems, 1902)

14. Anterior gonopods with small telopodital rudiments placed parabasally on lateral arms.

. A. albanensis Mauries, Golovatch & Stoev, 1997

- Anterior gonopods without telopodital rudiments.15

15. Position of ex-telopodites of posterior gonopods indicated by dark spots basally on angiocoxites

.16

- Angiocoxites of posterior gonopods without any trace of ex-telopodites.17

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European Journal of Taxonomy 227: 1-19 (2016)

16. Angiocoxites of posterior gonopods with 2-3 setae apically. A. pentelicona (Verhoeff, 1925)

- Angiocoxites of posterior gonopods serrated apically, but without setae.

. A. hauseri (Strasser, 1974)

17. Median part of anterior gonopods represented by a single, trapezoid plate ..A, alba (Strasser, 1960)

- Median part of anterior gonopods distally divided in two symmetric parts.18

18. Both colpocoxites and median part of anterior gonopods setose/spinulate at margins.

. A. falcata (Gulicka, 1967)

- Colpocoxites and median part of anterior gonopods without setiform or spiniform filaments.19

19. Angiocoxites of posterior gonopods about twice as long as colpocoxites, parallel to converging in

their proximal parts. A. lepenicae (Strasser, 1975)

- Angiocoxites of posterior gonopods less than twice as long as colpocoxites, diverging right from

their bases. A. delcevi (Strasser, 1973)

Discussion

Anamastigona terraesanctae, A. cypria sp. nov. and A. strasseri sp. nov. appear to be a natural group

characterized by the above described diagnostic features, as well as by generally similar morphometries

and colour pattern. This is yet another example of the considerable similarity between the millipede

faunas of Cyprus and the Levant (Vagalinski et al. 2014). Interestingly, A. strasseri sp. nov. seems

to occupy a morphologically intermediate position between A. terraesanctae and A. cypria sp. nov.,

sharing the uniramous lateral arms of the anterior gonopods and the stout, apically broad angiocoxites

of the posterior gonopods with the former species, but the very long and dense pilosity on the median

lamella and the elongated colpocoxites of the anterior gonopods with the latter.

At present, the exact number of species within Anamastigona remains uncertain. As already mentioned

in the introduction, the proposed synonymy of A. aspromontis (Strasser, 1970) with A. meridionalis

Silvestri, 1898 does not seem to be solid enough. There are a number of differences concerning the

gonopodal apparatus that can hardly be considered as mere individual variations: the anterior gonopods

in A. aspromontis have a more narrow median plate/lamella and bear two slender, apically branched

processes positioned between the colpocoxites and the lamella, these being absent in A. meridionalis ;

and, most importantly, the posterior gonopods in A. meridionalis possess very small angiocoxites, about

the same size as the telopodital rudiments, the former bearing several setae, vs the angiocoxites by

far outreaching the telopodital rudiments and being devoid of setae in A. aspromontis. Moreover, the

r

presence/absence of setae on the angiocoxites is considered an important diagnostic character by Curcic

et al. (2008), since these authors used it in their identification key in several couplets concerning the

species of Anamastigona. As regards A. halophila , it is not recognized as a valid species in the present

study, although the unknown conformation of the posterior gonopods in the single known male leaves a

certain amount of uncertainty about its possible synonymy with A bilselii (Verhoeff, 1940).

The inner grouping of Anamastigona is currently almost completely unresolved, with the only

recent consideration on the topic being that by Makarov et al. (2007), who divided the genus into

two groups based on the presence/absence of telopodites on the posterior gonopods and the shape of

their angiocoxites. According to their study, the Bulgarian endemics A. falcata (Gulicka, 1967), A. alba

(Strasser, 1960), A. lepenicae (Strasser, 1975) and A. delcevi (Strasser, 1973), possibly together with the

Albanian A. albanensis Mauries, Golovatch & Stoev, 1997, form a species group separate from the rest

of their congeners. However, in such a division A. hauseri (Strasser, 1974) and A. pentelicona (Verhoeff,

1925) should also join the first group, since their telopodites are completely reduced, with only some

pigmented spots/granules indicating the position of the lost appendages, like the condition observed in

14

Table 2. Main gonopodal characters of species of Anamastigona Silvestri, 1898.

VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

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15

European Journal of Taxonomy 227: 1-19 (2016)

A. albanensis. On the other hand, unlike the Bulgarian species, which possess very slender, hornlike

angiocoxites, the posterior gonopods in A. albanensis , A. hauseri and A. pentelicona are considerably

less-strongly modified. A strongly elongated shape of the angiocoxites is also present in A. bilselii

(Verhoeff, 1940) and A. radmani Makarov et al., 2007, although they bear well-pronounced telopodital

rudiments. The several main diagnostic gonopodal characters applied in the key by Curcic et al.

(2008) and in the present one are of little help for any clear intrageneric division to be proposed. Most

of the presumable derived characters (e.g., absence of telopodital rudiments on posterior gonopods

or absence of setae on angiocoxites of anterior gonopods) are reductive in nature and thus likely to

present homoplasies, rather than synapomorphies (see Table 2). A further difficulty is the presence of

unique, probably plesiomorphic characters in certain species, like the anterior gonopods possessing

flagella in A. matsakisi or telopodital rudiments in A. albanensis , both of these traits making it hard to

compare them to the remaining congeners. Perhaps the most outstanding species of all is A. mauriesr.

it displays a unique conformation of the posterior gonopods, which have unusually large colpocoxites

and telopodital rudiments placed on top of the angiocoxites, the latter condition supposed to represent

the most plesiomorphic state within the entire genus (Golovatch & Makarov 2011). On the other hand,

the considerably enlarged angiocoxites in combination with the strongly reduced, knot-like telopodites

may be the result of a different course of modification of the posterior gonopods that might have taken

place very early in the evolution of the genus. It is possible that future examination of vulval characters,

which with very few exceptions are completely unknown in Anamastigona, could shed more light on the

phylogenetic affinities of its constituent species.

Anamastigona is surely more speciose than we presently know. This can be assumed from the

small size of the animals, combined with their supposed low surface activity, as well as from the

overwhelmingly local endemism. Vast, insufficiently explored areas like the Mediterranean coast of

Turkey and, especially, the Caucasus, the latter considered as a diversity hotspot for the entire family

Anthroleucosomatidae (Mauries et al. 1997; Golovatch & Makarov 2011), are probably inhabited by a

number of still undescribed members of the genus.

Checklist of Anamastigona species

Anamastigona alba (Strasser, 1960)

Distribution

Bulgaria (Enghoff & Kime 2013).

Anamastigona albanensis Mauries, Golovatch & Stoev, 1997

Distribution

Albania (Enghoff & Kime 2013).

Anamastigona aspromontis (Strasser, 1970)

Distribution

Italian mainland (Enghoff & Kime 2013).

Anamastigona bilselii (Verhoeff, 1940)

Distribution

Bulgaria (Strasser 1973), European Turkey, Greek mainland (Enghoff & Kime 2013).

Anamastigona cypria sp. nov.

Distribution

Cyprus.

16

VAGALINSKIB. & GOLOVATCH S.I., Two new species of Anamastigona from Cyprus

Anamastigona delcevi (Strasser, 1973)

Distribution

Bulgaria (Enghoff & Kime 2013).

Anamastigona falcata (Gulicka, 1967)

Distribution

Bulgaria (Enghoff & Kime 2013).

Anamastigona hauseri (Strasser, 1974)

Distribution

Greek mainland (Enghoff & Kime 2013).

Anamastigona hispidula (Silvestri, 1895)

A. napolitana (Attems, 1899)

A. apenninorum (Verhoefif, 1930)

A. hispidula ischiana (Verhoeff, 1940)

A. hiemalis (Verhoeff, 1941)

A. hispidula quercuum (Verhoeff, 1942)

A. hispidula relicta (Verhoeff, 1942)

A. hispidula epome ana (Verhoeff 1952)

Distribution

Italy: mainland (Enghoff & Kime 2013), Capri (Verhoeff 1942), Ischia (Verhoeff 1940a, 1941).

Anamastigona lepenicae (Strasser, 1975)

Distribution

Bulgaria (Enghoff & Kime 2013).

Anamastigona matsakisi Mauries & Karamaouna, 1984

Distribution

Greece: Naxos (Mauries & Karamaouna, 1984).

Anamastigona mauriesi Golovatch & Makarov, 2011

Distribution

Autonomous Republic of Abkhazia (Golovatch & Makarov 2011).

r

Anamastigona mediterranea Curcic, Makarov & Lymberakis, 2001

Distribution

Greece: Crete (Enghoff & Kime 2013).

Anamastigona meridionalis Silvestri, 1898

Distribution

Italy: Sicily (Enghoff & Kim e 2013).

Anamastigona penicillata (Attems, 1902)

Distribution

Greece: Crete (Enghoff & Kime 2013).

Anamastigona pentelicona (Verhoeff, 1925)

Distribution

Greek mainland (Enghoff & K im e 2013).

17

European Journal of Taxonomy 227: 1-19 (2016)

Anamastigona pulchella (Silvestri, 1894)

A. attemsi (Verhoeff, 1900)

A. macchiae (Verhoeff, 1930)

Distribution

Italian mainland, French mainland, Madeira, Northern Ireland (Enghoff & Kime 2013), Germany

(Lindner et al. 2010).

Anamastigona radmani Makarov et al. , 2007

Distribution

Croatia (Makarov et al. 2007).

Anamastigona strasseri sp. nov.

Distribution

Cyprus.

Anamastigona terraesanctae Golovatch & Makarov, 2011

Distribution

Israel (Golovatch & Makarov 2011).

Acknowledgements

We are most thankful to Stylianos Simaiakis (NHMC) and to Kirill Mikhailov (ZMUM) for making

specimens under their care available for study. Yordan Keremidchiev (Sofia, Bulgaria) skillfully refined

the SEM- and stereo microscopic pictures.

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Manuscript received: 29 January 2016

Manuscript accepted: 6 April 2016

Published on: 6 September 2016

Topic editor: Rudy Jocque

Desk editor: Danny Eibye-Jacobsen

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands.

19