European Journal of Taxonomy 70: 1-12

http://dx.doi.org/lQ.5852/ejt.2Q13.7Q

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

WWW, europeanj oumaloftaxonomy. eu

2013 • Lazanyi E. & Vagalinski B.

Research article

urn:lsid:zoobank.org:pub:5E23E454-2A68-42E6-86EB-7D9952B2EE7B

A new genus of the millipede tribe Brachyiulini (Diplopoda:

Julida: Julidae) from the Aegean region

Eszter LAZANYI'-'' & Boyan VAGALINSKP-^-^

^ Corresponding author: Department of Zoology, Hungarian Natural History Museum, Baross u. 13,

H-1088 Budapest, Hungary. E-mail: [email protected]

^ Faeulty of Biology, Sofia University, 8 Dragan Tsankov Blvd., 1164 Sofia, Bulgaria.

E-mail: [email protected]

^ Institute of Biodiversity and Eeosystem Researeh, Bulgarian Aeademy of Seienees, 2 Yurii Gagarin

Street, 1113, Sofia, Bulgaria.

um:lsid:zoobank.org:author:Q2DB48Fl-624C-4435-AF85-FA87168CD85A

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

Abstract. Anew genus of the julid tribe Braehyiulini, Enghophyllum gen. nov., is deseribed, eomprising

two speeies from Greeee. The type-speeies, E. naxium (Verhoeff, 19Q1) eomb. nov. (ex Megaphyllum

Verhoeff, 1894), appears to be rather widespread in the Aegean: it is known from Antiparos Island

and Naxos Island (the type loeality), both in the Cyelades, as well as East Mavri Islet, Dodeeanese

Arehipelago (new reeord). The vulva of E. naxium is deseribed for the first time. In addition, E. sifnium

gen. et sp. nov. is deseribed based on a single adult male from Sifnos Island, Cyelades. The new genus

is distinet from other genera of the Braehyiulini mainly by its peeuliar gonopod strueture, apparently

disjunet and at least mostly apomorphous: (1) promeres broad, shield-like, in situ protruding mostly

posteriad, eompletely eovering the opisthomeres and gonopodal sinus; (2) transverse museles and

eoxal apodemes of promere fully redueed; (3) opisthomere with three differentiated proeesses, i.e.,

lateral, basal posterior and apieal posterior; (4) solenomere rather simple, tubular. The evolution and

biogeography of the new genus are briefiy diseussed, both suggesting its profoundly long isolation in

the Aegean region from the eontribal genera in the adjaeent Balkans and Anatolia.

Key words. Brachyiulus, Chromatoiulus, Megaphyllum, new taxa, new reeords.

Lazanyi E & Vagalinski B. 2Q13. A new genus of the millipede tribe Brachyiulini (Diplopoda: Julida: Julidae) from

the Aegean region. European Journal of Taxonomy 7Q: 1-12. http://dx.doi.org/lQ.5852/eit.2Q13.7Q

Introduction

The present paper eontinues our efforts in revising the large julid tribe Braehyiulini (Fazanyi et al. 2Q12;

Fazanyi & Vagalinski 2Q13; Vagalinski et al. 2Q13), this time being devoted to the deseription of a new

oligotypie genus from several Aegean islands of Greeee. Its type speeies, E. naxium (Verhoeff, I9QI),

previously referred to as Brachyiulus naxius, Chromatoiulus (Diaxylus) naxius or Megaphyllum naxium,

has been found to oeeur both in the Cyelades and Dodeeanese arehipelagos, whereas the seeond eongener,

E. sifnium gen. et sp. nov., is a new speeies deseribed below. The gonopod and, to some degree, the

European Journal of Taxonomy 70 : 1-12 ( 2013 )

vulval structures of these two speeies appear to be of great interest by shedding additional light on the

evolution and biogeography of the entire fauna of eastern Mediterranean Braehyiulini.

Material and methods

The present work is based on the examination of material from the Non-inseet Invertebrate Colleetion,

National Museum of Natural History (NMNHS), Sofia, and the Natural History Museum of Denmark,

Zoologieal Museum, University of Copenhagen (ZMUC). Types of the new speeies are deposited in

ZMUC. Speeimens are stored in 70% ethanol and small disseeted parts have been plaeed in genitalia

vials whieh are kept together with the disseeted individuals. Female vulvae were mounted on slides in

Faure-Berlese medium and ineubated at room temperature (around 25°C) for several hours. After having

been studied, the slides were put into distilled water, vulvae were removed from the Faure-Berlese

medium and put into 70% ethanol.

Line drawings were made with a Leiea Ml25 (HNHM) stereo mieroseope and an Olympus BX40 light

mieroseope.

The apieal end of the opisthomere is distinguished as the solenomere; all proeesses emerging from

the opisthomere’s main body are named simply aeeording to their position as lateral, basal and apieal

posterior proeesses.

Results

Class Diplopoda Blainville-Gervais, 1844

Order Julida Leaeh, 1814

Family Julidae Leaeh, 1814

Tribe Braehyiulini Verhoeff, 1909

Genus Enghophyllum gen. nov.

um:lsid:zoobank.org:aet:15008961-CD77-4325-B694-13BAlF7E514E

Type species

Brachyiulus (Chromatoiulus) naxius Verhoeff, 1901

Diagnosis

A genus of Braehyiulini, differing from eontribal genera by the following eombination of eharaeters:

promere broad, shield-like, in situ protruding mostly posteriad, eompletely eovering opisthomere and

gonopodal sinus; transverse museles and eoxal apodemes of the promere fully redueed. Opisthomere

with 3 well-differentiated proeesses: lateral {Ip), basal posterior {bpp) and apieal posterior {app).

Solenomere rather simple, tubular.

Etymology

This genus is named in honor of Prof Hemik Enghoff from the ZMUC, not only for his vast eontribution

to our knowledge of diplopods, but also for his eontinuous eneouragement and help during our work

with Braehyiulini.

Description

Two frontal, four supralabral and 14-18 labral setae. Antennomeres 4 and 5 subequal in length (Fig. lA).

Gnathoehilarium as on Fig. IB. Metazona sparsely striated, as emphasised by Verhoeff (1901); ozopores

right on the suture (Fig. 1C). Preanal proeess straight, anal valves sparsely pilose (Fig. ID); subanal

seale triangular, with protruding tip (Fig. IE).

LAZANYIE. & VAGALINSKIB., A new genus of Braehyiulini from the Aegean region

Males

Mandibular stipes slightly protruding (Fig. IF-G). First leg-pair like simple, rounded, somewhat

eonverging hooks; the two hooks eonverging in an obtuse angle (Fig. IH). Seeond leg-pair (Fig. II) with

two ventral pads: on postfemur {pf) and tibia (/). Penis (Fig. IJ) signifieantly small, deeply hidden in

penis sae, with two very short lobes and two rounded small lamellae running parallel, i.e., not diverging

(r/ and ll for right and left lamella). Pleurotergum of the body ring protruding like a simple shovel

(broken in the unique holotype of E. sifnium gen. et sp. nov., thus investigated only in E. naxium, see Fig.

Fig.l . Non-gonopod eharaeters ofEnghopyhllum gen. nov., males. A. Right antenna. B. Gnathoehilarium.

C. 8* and 9* body rings. D. Telson. E. Subanal seale. F. Mandibular stipital (“eheek”) lobe, lateral view.

G. Mandibular stipital (“eheek”) lobe, ventro-lateral view. H. First leg-pair. I. Seeond leg-pair. J. Penis

in the penis sae. K. Pleurotergum of the 7* body ring, ventral view. L. Anterior left leg on the 6* body

ring. M. Anterior right leg on the 22"^ body ring. - A-D, F, H-I, F-M: E. sifnium gen. et sp. nov., holotype.

E: E. naxium, from Antiparos. G, J-K: E. naxium, from Mavri Islet.

European Journal of Taxonomy 70 : 1-12 ( 2013 )

IK). Walking legs (Fig. IL-M) with two ventral pads: on postfemur ipf) and tibia (/). Tarsus of midbody

legs slightly longer than tibia: tarsus / tibia = (1.05-1.1) / 1.

Gonopods notieeably big eompared to body size, direeted mostly posteriad. Promere bevelled in latero-

basal eomer to fit into the inner eurve of the 7* body ring; latero-basal eomer of promere only with

thin triangular lamella {tl) (Fig. 2D-E). Promeres attaehed to eaeh other in their meso-basal eomer with

a ehitinous bridge. A short, oblique ridge (r) on posterior surfaee of promere (Figs 2A, D-E, 4A, C).

Flagellum emerging from a well-developed sinus ifsi) (Fig. 2D-E). At the level of this sinus a deepening

on anterior surfaee of the promere {d) (Fig. 4B). Opisthomere entirely eoneealed under the promere,

protmding postero-ventrad, the two promeres tightly elosing the ventral opening on 7* ring like a pair

of shutters. Opisthomere not embedded in the groove formed by the oblique ridge on the promere’s

posterior surfaee (unlike in Megaphyllum), but “riding” on the ridge (Figs 2A, 4C): posterior basis of

opisthomere with a eharaeteristie riffle (ri) formed by ehitinous lamellae, where the ridge of the promere

is positioned (Figs 2A, G, 4C-D). Opisthomere with a well-developed lateral proeess (Ip) and an apieal

and a basal posterior proeess (app and bpp). Solenomere (s) simple, with no proeesses but basally with

a miero-spinose pillow (msp) (Figs 2A-C, F-G, 4A-D). Flagellum ehannel and sperm eanal (fc and sc)

eaeh with an apieal opening (Fig. 4C).

Females

The only available female of E. sifnium gen. et sp. nov. is in stadium IX (with 8 rows of oeelli) and

proved to be subadult, but females of E. naxium in stadia Vlll-lX (with 7-8 rows of oeelli) had fully

developed vulvae, whieh we used here for the deseription of female sexual eharaeters. First two leg-

pairs slightly swollen.

Vulva (Fig. 3): Subeylindrie in shape, the mesal half shorter, the opening large, oval, apieal. Opereulum

shorter than bursa, with around 20-25 setae. Mesal and lateral selerites with 2-3 setae eaeh. Apodematie

tube not opening apieally into a longitudinal median eleft as in, e.g., Megaphyllum speeies, but into a

wide sinus (si). Around this sinus a thin wall, formed by the apieal part of the bursa. These walls with

small dot-like pores; most setae (around 20/side) emerging from this apieal region. Apodematie tube (at)

ending in two reeeptaeuli seminis or ampullae. Central ampulla (ca) drop-like. Distal, globular ampulla

((ia) joining the apodematie tube through a twisted eonneeting tube (ct). Conneeting tube very long, thus

the distal ampulla hanging out of the bursa; easily broken off during preparation.

Remarks

The everted penes of the adult E. naxium male from Antiparos was quite soft, amorphous and huge,

eontrary to the stout, minute penes, deeply hidden under the 2"^ leg-pair eoxae, observed in the E.

naxium male from Mavri Islet and the E. sifnium gen. et sp. nov. male from Sifnos. It is possible that

penes vary in size between eopulating and non-eopulating periods.

Enghophyllum sifnium gen. et sp. nov.

um:lsid:zoobank.org:aet:5327BB09-7438-47BD-9D91-BBC6FE7FlCC5

Figs lA-D, F, H-1, E-M, 2A-G, 5

Diagnosis

Differs from the only eongener, E. naxium (Verhoeff, 1903), by the apieal margin of the promere being

slightly eonvex and the opishtomere being signifieantly eompressed meso-laterad.

Etymology

Named after the type loeality, the island of Sifnos, Cyelades.

LAZANYIE. & VAGALINSKIB., A new genus of Braehyiulini from the Aegean region

Material examined

Holotype

GREECE: S, Hellas [Greeee], Kildadhes [Cyelades], Sifnos Island, Pharos [Faros], ll.ii.l990 [11 Feb.

1990], S. Sfendourakis leg., ex eoll. Karamaouna, ''Megaphyllum aff. naxium (Verhoeff, 1901)” M.

Karamouna det. (ZMUC).

Fig. 2. Gonopods ofEnghophyllum sifnium gen. et sp. nov., holotype S- A. Eeft gonopods, mesal view. B.

Eeftgonopods, antero-lateralview. C. Eeftgonopods,meso-posteriorview. D. Eeftpromere,meso-posterior

view. E. Eeft promere, postero-mesal view. F. Right opisthomere, lateral view. G. Right opisthomere,

mesal view. - app = apieal posterior proeess; bpp = basal posterior proeess; fc = flagellum ehannel;

fl = flagellum; fsi = flagellum sinus; Ip = lateral proeess; msp = miero-spinose pillow; P = promere;

r = ridge; ri = rifiie; ^ = solenomere; tl = triangular lamella. Seale bar: 0.2 mm.

European Journal of Taxonomy 70 : 1-12 ( 2013 )

Paratypes

GREECE: 1 juv. S, 1 juv. $. Same collection data as holotype (ZMUC).

Description

Holotype male in stadium X (with 9 rows of ocelli); body ring number: 41 podous + 2 apodous + telson;

length 18.8 mm; height 1.1 mm; ocelli: right side: 35, left side: 33. Additional measurements of the adult

male: promentum around twice as long as wide (Fig. IB), tibia of midbody legs slightly longer than

tarsus, 4^*^ antennal article as long as 5^*^ (Fig. lA). Subadult paratype male in stadium IX (with 8 rows

of ocelli); body ring number: 38 podous + 3 apodous + telson; length 13.1 mm; height 1.1 mm; ocelli:

right side: 34, left side: 35. Subadult female in stadium IX (with 8 rows of ocelli); body ring number: 39

podous + 3 apodous + telson; length 14.1 mm; height 1 mm; ocelli: right side: 30, left side: 30. 17-18

labral bristles.

Colour. Basic colour yellowish-beige (obviously faded) with a narrow brownish middorsal band,

divided by a white median line. Two wider, diffused brownish bands at ozopore level, more intensely

coloured around the ozopores.

Sculpture. Collum smooth, with only several short striae visible; metazona sparsely striated (Fig. 1C),

somewhat denser on dorsal side, posterior margin with long, erect setae. Scarce, short striae also on

prozona.

Telson (Fig. ID). Anal valves covered with scarce, long setae and a row of much shorter setae along

each valve’s caudal margin. Preanal process without any setae in the adult male, but with a few setae

in the two subadults; straight, stout, protruding near to the level of the longest anal setae (for the adult

male). Subanal scale triangular, with a rounded tip, slightly protruding behind rear contour of anal

valves; with two pairs of long setae laterally.

Gonopods (Fig. 2A-G). Promere (Fig. 2D-E and P on Fig. 2A-C) wide, slightly narrowing apically

towards mesal margin. Apical margin convex, mesal and lateral margins almost parallel, the lateral

one slightly sinuous. Opisthomere with three processes: a flat lateral process {Ip on Fig. 2A-C, F), a

slightly curved, rod-like basal posterior process {bpp on Fig. 2A-C, F-G) and an elongate, massive,

rod-like apical posterior process {app on Fig. 2A-C, F-G); all processes well-developed, tapering and

not furcated. A micro-spinose pillow {msp) around the base of solenomere {s) (Fig. 2A-C, F-G). Entire

opisthomere strongly compressed meso-laterad (compare Fig. 2C with Fig. 4A).

Female adults unknown.

Enghophyllum naxium (Verhoeff, 1901) comb. nov.

Figs IE, G, J-K, 3, 4A-D, 5

Brachyiulus (Chromatoiulus) naxius Verhoeff, 1901: 98-99, flg. 21.

Chromatoiulus naxius - AttQms 1921 \ 222 (in key). — Strasser 1974: 290, 295. — Ceuca 1992: 425.

Chromatoiulus (Diaxylus) naxius - AttQms 1940: 307, 311-312.

Megaphyllum naxium - Mauries & Karamaouna 1984: 55-56, flgs 7-9. — Enghofif & Kime 2009. —

Eazanyi et al. 2012: 21,41.

Diagnosis

Differs from the only congener E. sifnium gen. et sp. nov. by the apical margin of the promere being

concave, the opisthomere not compressed meso-laterad, e.g., the lateral process {Ip on Fig. 4A-D)

parallel to the promere, the lateral process {Ip) much broader and the apical posterior process {app)

broader and blunter.

LAZANYIE. & VAGALINSKIB., A new genus of Braehyiulini from the Aegean region

Material examined

GREECE: 1 (J, 2 Antiparos Island, litiere [litter], 22 Dee. 1982, leg. R Beron & St. Andreev,

det. Mauries 1983 (NMNHS); 1 Sporadhes, Anatoliko Mauri [east Mavri Islet] (Eevitha Islands),

18.i. 1990 [18 Jan. 1990], S. Sfendourakis leg., ex eoll. M. Karamaouna (ZMUC).

Remarks

Male body length 16.9-17.9 mm; height 1-1.2 mm; body ring number: 39-41 podous rings + 2-3 apodous

rings + telson; number of oeelli: right side: 30-38, left side: 32-38; developmental stadium IX-X with

Fig. 3. Right vulva of Enghophyllum naxium, female from Antiparos Island, anterior view. -

at = apodematie tube; ca = eentral ampulla; ct = eonneeting tube; da = distal ampulla; Iv = lateral valve;

op = opereulum; si = sinus. Seale bar: 0.2 mm.

European Journal of Taxonomy 70 : 1-12 ( 2013 )

8-9 rows of ocelli. Female body length 13.4-18.3 mm; height 1-1.3 mm; body ring number: 37-41

podous rings + 3-5 apodous rings + telson; developmental stadium Vlll-lX with 7-8 rows of oeelli.

This speeies was deseribed on the basis of two males and four females and has only been found onee

sinee its deseription, viz., by Mauries & Karamaouna (1984) from Antiparos. The new speeimens from

east Mavri belong to the eolleetion of Karamaouna, but they are slightly different: the apieal posterior

proeess (app) of the solenomere is not bifureated, but undivided and slightly serrated (serration visible

in posterior view) (Fig. 4A, C-D).

Fig. 4. Gonopods of Enghophyllum naxium, male from Mavri Islet. A. Gonopods, posterior view.

B. Gonopods, antero-lateral view. C. Right gonopods, mesal view. D. Left opisthomere, lateral view.

- app = apieal posterior proeess; bpp = basal posterior proeess; d = deepening; fc = flagellum ehannel;

Ip = lateral proeess; msp = miero-spinose pillow; P = promere; r = ridge; ri = riffle; ^ = solenomere;

sc = sperm eanal; tl = triangular lamella. Seale bar: 0.2 mm.

LAZANYIE. & VAGALINSKIB., A new genus of Braehyiulini from the Aegean region

Distribution

Greeee: Cyelades: Naxos Island (type loeality), Antiparos Island (Mauries & Karamaouna 1984);

Dodeeanese Islands: east Mavri Islet.

Discussion

Enghophyllum naxium was hitherto treated as a Megaphyllum speeies (or under its former synonyms

Chromatoiulus and Brachyiulus s. auct.) (Lazanyi et al. 2012). It was plaeed in the subgenus Diaxylus

Attems, 1940 (type speeies: Chromatoiulus anatolicus AttQms, 1927) on the basis of having a simple

promere, a simple solenomere, a posterior proeess at right angles to the opisthomere and laeking a

bristle-row along the flagellum ehannel (Attems 1940). After thorough investigations of this speeies and

ofE. sifnium gen. et sp. nov., supplemented with the examination of Megaphyllum argolicum (Verhoeff,

1900), M. asiaeminoris (Verhoeff, 1898), M. euphorbiarum (Verhoeff, 1900) and M. anatolicum

(Attems, 1927), all referred to Diaxylus, it beeame apparent that E. naxium and E. sifnium gen. et sp.

nov. form a group neither with the aforementioned speeies, nor with any other Megaphyllum subgenus

or Braehyiulini genus. The eombination of external morphologieal eharaeters together with male and

female genitalia eharaeters supports the establishment of a new braehyiuline genus: Enghophyllum gen.

nov. Considering gonopodal strueture, some similarities with Megaphyllum byzantinum (plaeed in the

monotypie subgenus Byzantorhopalum Verhoeff, 1930) ean be diseemed, namely the well-developed

lateral proeess of the opisthomere and the eylindrieal vulvae with a mostly apieally positioned opening.

However, some external and internal features of the vulvae, the shape of the penes, and notably the

unique arrangement of the promeres leave no doubt that these speeies belong to a different genus.

Fig. 5. Distribution map of Enghophyllum gen. nov. The eheeked area (Silhos Island) represents the

possible distribution of E. sifnium gen. et sp. nov.; the red star shows Faros, its type loeality. The

obliquely lined areas (Antiparos and Naxos Islands, Mavri Islet) show the distribution of E. naxium.

European Journal of Taxonomy 70 : 1-12 ( 2013 )

The known distribution of Enghophyllum gen. nov. is so far restrieted to three islands in the Cyelades

(Naxos, Antiparos and Sifnos) and one small islet in the westernmost part of the Dodeeanese Islands,

Greeee (east Mavri Islet, Fig. 5). There are eurrently 10 other speeies of Braehyiulini reported from

the Aegean Islands: Brachyiulus stuxbergi (Fanzago, 1875) (Crete), Megaphyllum bicolor (Loksa,

1970) (Rhodos, Naxos), M. brachyurum (Attems, 1899) (Thasos), M. chiosense Lazanyi & Korsos,

2012 (Chios), M. creticum (Strasser, 1976) (Crete), M. mueggenburgi (Verhoefif, 1901) (Karpathos,

Kasos), M. rossicum (Timotheew, 1897) (Samothraki), M. sapphicum (Strasser, 1976) (Lesvos),

M. syrense (Verhoeff, 1903) (Syros) mdM. taygeti (Strasser, 1976) (Crete) (Felesaki etal. 2010; Lazanyi

et al. 2012). Most of them are known only from a single island, exeept for M. mueggenburgi, whieh

inhabits the adjaeent Karpathos and Kasos, and M. bicolor with its nominotypieal subspeeies deseribed

from Rhodos and the subspeeies M. b. crassiflagellum deseribed from Naxos. It is also notable that

only 4 speeies, namely B. stuxbergi, M. brachyurum, M. taygeti and M rossicum, oeeur both on islands

and on either European or Asian mainlands. B. stuxbergi is widely distributed in Greeee and Italy; M.

brachyurum is represented on Thasos with its subspeeies M. b. thassensis, while the nominotypieal

subspeeies inhabits the Caueasian region; M. taygeti oeeurs on the Peloponnese; M. rossicum rossicum

is widely distributed in South Russia and the Ukraine, while M. r strandschanum has been reported from

south Bulgaria and northeast eontinental Greeee, apart from the single reeord from Samothraki.

The Aegean Islands seem to support a high number of endemies of the Braehyiulini tribe, the most

outstanding of whieh are the two speeies of Enghophyllum gen. nov. Compared to a number of

biogeographie and phylogeographie papers dealing with different invertebrate groups from the Aegean

region (e.g., Douris et al 2007; Fattorini & Fawles 2005; Sfenthourakis 1996; Welter-Sehultes &

Williams 1999) the above deseribed distribution pattern of the Aegean Braehyiulini is espeeially similar

to that observed by Heller (1976). He found most Aegean speeies of the terrestrial gastropod family

Enidae to oeeur only on one or several adjaeent islands, whieh is eonsidered to refleet the laek of aetive

immigration in reeent geologieal times. These biogeographie similarities eould be explained with the

very limited dispersal abilities eharaeteristie of millipedes and land snails in general. Another reason

may be the rather weak eeologieal speeialization eommon to both groups. Thus, their eurrent distribution

would be less affeeted by the presenee/absenee of partieular plant eommunities or habitat degradation,

unlike the ease with, e.g., the eentral Aegean Oniseidea (Sfenthourakis 1996), and would more elearly

refleet the geologieal events that took plaee in the Aegean region. In this respeet the reeord for E. naxium

from east Mavri may present interesting information in terms of further revealing the biogeographie

affiliations of this group of several small islands (Kinaros, west and east Mavri and Eevitha), positioned

between the Cyelades and the Dodeeanese although ofheially aseribed to the latter. Sfenthourakis (1996)

suggested that Mavri is biogeographieally related to several seattered islands in the south Cyelades,

while Eevitha has more in co mm on with the other Dodeeanese islands. However, the overal searee

faunistie data hamper any Arm eonelusions being reaehed about the islets’ paleogeographie history.

In spite of the rather few distribution reeords for the new genus, as well as the insuffleiently investigated

millipede fauna of the Greek islands in general, we may suppose that the genus deseribed here represents

a distinet lineage of Braehyiulini that has developed in relatively long-lasting isolation, as there are

eurrently no apparent elosely related forms in the Balkans or in Anatolia. The reliet eharaeter of the

Aegean fauna has already been demonstrated for other invertebrate groups like tenebrionid beetles

(Fattorini & Fawles 2005) and land snails (Heller 1976; Welter-Sehultes & Williams 1999).

In eonelusion, it ean be assumed that Enghophyllum gen. nov. eonflrms the importanee of the Aegean

Islands together with south Greeee and west Turkey for the evolution of the millipede tribe Braehyiulini,

being inhabited by both highly distinet loeal forms, as well as by members of (sub)genera and speeies

groups found in other regions - evidenee of sueeessive proeesses of migration and viearianee resulting

LAZANYIE. & VAGALINSKIB., A new genus of Braehyiulini from the Aegean region

in intense speeiation. On the other hand, the dominant position of the tribe in terms of speeies ric hn ess

among Diplopoda from the region, eombined with a high rate of endemism, provide good reasons for

using the group as a model for further elueidation of the eomplieated paleogeographie history of the

Aegean Arehipelago.

Acknowledgements

We would like to thank Prof Hemik Enghoff at the Natural History Museum of Denmark and Prof

Pavel Stoev at the National Museum of Natural History, Bulgaria for their help and the arrangement of

loans of type and non-type speeimens.

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Manuscript received: 1 July 2013

Manuscript accepted: 8 November 2013

Published on: 17 December 2013

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 eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium; National History

Museum, Fondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.