European Journal of Taxonomy 195: 1—47

http://dx.doi.org/10.5852/ejt.2016.195

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2016 • Likhitrakarn N. et al.

Research article

urn:lsid:zoobank.org:pub:09F2E058-97B4-4651-B306-0975B4EE490F

The millipede genus Tylopus Jeekel, 1968 (Diplopoda, Polydesmida,

Paradoxosomatidae), with a key and descriptions

of eight new species from Indochina

Natdanai LIKHITRAKARN 1 , Sergei I. GOLOVATCH 2 * & Somsak PANHA 3 -*

’Division of Plant Protection, Faculty of Agricultural Production,

Maejo University, Chiang Mai 50290, Thailand,

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

Leninsky pr. 33, Moscow 119071, Russia.

3 Animal Systematics Research Unit, Department of Biology, Faculty of Science,

Chulalonglcorn University, Bangkok 10330, Thailand.

’Email: [email protected]

^Corresponding author: [email protected]

3 *Corresponding author: [email protected]

1 urn:lsid:zoobank.org:author:442E2E6E-lCB3-47B2-9896-lA922CE509El

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

3 um:lsid:zoobank.org:author:AC935098-D901-4F35-A414-4B0D4FE44E79

Abstract. The Indochinese to southern Chinese millipede genus Tylopus currently comprises 62 species,

including eight new ones: T. flavolineatus sp. nov., T. hongkhraiensis sp. nov. from central and northern

Thailand, respectively, T. moniliformis sp. nov., T. retusus sp. nov., T. acuminatus sp. nov., T. dorsalis

sp. nov., T. thunghaihin sp. nov., all from Laos, and T. punctus sp. nov. from northern Myanmar. Anew

record of T. baenzigeri Golovatch & Enghoff, 1993, from northern Thailand is given. All these species

are richly illustrated, and a modified key to all known species of the genus is also provided.

Keywords. Myriapoda, Taxonomy, Thailand, Laos, Myanmar.

Likhitrakarn N., Golovatch S.I. & Panha S. 2016. The millipede genus Tylopus Jeekel, 1968 (Diplopoda,

Polydesmida, Paradoxosomatidae), with a key and descriptions of eight new species from Indochina. European

Journal of Taxonomy 195: 1-47. http://dx.doi.org/10.5852/eit.2016.195

Introduction

Tylopus Jeekel, 1968, has long been recognized as one of the largest and widespread millipede Oriental

genera in the entire family Paradoxosomatidae (Likhitrakarn et al. 2010). This genus has been reviewed

and rediagnosed several times, chiefly in the context of adding new species (Jeekel 1965,1968; Golovatch

& Enghoff 1993; Likhitrakarn et al. 2010). One of the most recent updates considers and keys 41 species

(Likhitrakarn et al. 2010), followed by Nguyen (2012) who added five new congeners from Vietnam

and provided a key and a distribution map to all 13 Tylopus known to occur in that country. Liu & Luo

1

European Journal of Taxonomy 195 : 1-47 ( 2016 )

(2013), and Golovatch (2013, 2014) described a few other new Tylopus species from southern China.

The latest review of Tylopus in Thailand (Likhitrakarn et al. 2014a) provides a new distribution map and

an updated key to all 32 species known to occur in Thailand. As a result, Tylopus is currently recognized

as the largest genus of Diplopoda in the Oriental realm, comprising 54 described species ranging from

southern China (6) to Myanmar (2), Taos (1), Thailand (32) and Vietnam (18).

Almost all Tylopus species appear to be confined to high montane forest habitats (Likhitrakarn et al.

2014a), usually representing highly local endemics. This genus is readily distinguished from all

Sulciferini (Paradoxosomatidae) except Oxidus Cook, 1911, by gonopodal postfemoral lamina 1 being

separated from the femorite by a more or less distinct transverse sulcus, from the oligotypic Oxidus

by the number and location of postfemoral outgrowths or processes (Golovatch & Enghoff 1993). In

addition, most of Tylopus species show adenostyles (= ventral outgrowths) at least on some of the male

legs. Only very few species, e.g., T. peramatus Hoffman, 1973, which is quite widespread in northern

Thailand (Hoffman 1973; Likhitrakarn etal. 2010), demonstrate evident variations in gonopod structure,

whereas it is gonopod conformation that provides many of the key characters for species identification.

Numerous Tylopus species are found to occur sympatrically in such larger montane areas as Doi Inthanon

(10 species) and Doi Suthep national parks (again 10 species, mostly different) (Likhitrakarn et al. 2010,

2014a) in Thailand, but the adult, identifiable stages of at least some of them seem to represent distinct

phenofaunas restricted to rather short time periods which fail to overlap with others.

The present paper puts on record another eight new species of Tylopus , this time from Myanmar, Thailand

and Laos, and their distributions are mapped. A new, modified key to all 62 species of the genus is also

given.

Material and methods

New material was collected from northern Thailand and southern Laos from 2008 to 2014 by SP

and members of the Anim al Systematics Research Unit, Chulalonglcorn University. Live an im als

were photographed in the laboratory. Specimens were preserved in 75% ethanol, and morphological

investigations were carried out in the laboratory using an Olympus stereo microscope. Scanning electron

micrographs (SEM) of gonopods coated with gold were taken using a JEOL, JSM-5410 LV microscope,

and returned to alcohol after examination to ultimately be stored in the alcohol collection. Digital images

of the specimens were taken in the laboratory and assembled using the “Cell 0 ” automontage software

of the Olympus Soft Imaging Solution GmbH package. In addition, line drawings of gonopods were

also prepared. All holotypes, as well as most of the paratypes, are kept in the Museum of Zoology,

Chulalonglcorn University (CUMZ), Bangkok, Thailand; some duplicates were also donated to the

collections of the Natural History Museum of Denmark, University of Copenhagen, Denmark (ZMUC),

the Zoological Museum, State University of Moscow, Russia (ZMUM), and the Naturhistorisches

Museum Wien, Austria (NHMW), as indicated in the text. The types of one species from Myanmar were

revised using the NHMW collection.

Collecting sites were located by GPS using the WGS84 datum.

The terminology concerning gonopod structure and body characters mostly follows Golovatch &

Enghoff (1993) and Likhitrakarn et al. (2010).

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LIKHITRAKARN N. et al. , The millipede genus Tylopus

Results

Taxonomic descriptions

Class Diplopoda Blainville-Gervais, 1844

Order Polydesmida Leach, 1814

Family Paradoxosomatidae Daday, 1889

Subfamily Paradoxosomatidae Daday, 1889

Tribe Sulciferini Attems, 1898

Genus Tylopus Jeekel, 1968

Tylopus flavolineatus sp. nov.

urn:lsid:zoobank.org:act:4A4DA74B-774B-45F5-A60F-96EB516022D2

Figs 1-3, 26

Diagnosis

Differs in the presence of middle and lateral yellow stripes on the body, by the sterna bearing evident

cones caudally near each coxa, and in $ femur 5 showing a large distovental adenostyle, coupled with

the gonopods being rather slender, with a quite large, slightly curved and pointed process h.

Etymology

To emphasize the middle and lateral yellow stripes on the body; adjective.

Material examined

Holotype

THAILAND: S, Wat Tham Santisulc, Nong Muang, Lopburi, 15°12T0" N, 100°39'50" E, 145 m a.s.l.,

7 Jun. 2008, leg. C. Sutcharit & N. Likhitrakarn (CUMZ).

Paratypes

THAILAND: 2 <$<$, 2 $ $ (CUMZ), same data as for holotype.

Description

Measurements and colour. Length 13.8-16.4 (c?) or 14.2-16.5 mm (J), width of midbody pro- and

metazonae 0.83-0.92 and 1.21-1.33 mm (c?) or 1.19-1.28 and 1.68-1.73 mm (§), respectively. Coloration

of alcohol material after six years of preservation dark brown; a pair of paramedian longitudinal stripes

flanking a narrow, sometimes vague, brown, axial line; paraterga and area of pleurosternal carinae

contrasting light yellow to yellowish; legs and venter dark brown to dark yellow (Fig. 1).

Head. Clypeolabral region densely, vertex sparsely, setose; epicranial suture distinct. Antennae rather

short (Fig. IB), clavate (antennomere 6 highest), extending behind body segment 3 (c?) or 2 (§) when

stretched dorsally. In width, segment 2 = 3 < head < collum < segment 4 < 5-15; thereafter body

gently and gradually tapering. Collum with three transverse rows of very small setigerous tubercles:

6+6 anterior, 4+4 intermediate and 5+5 posterior; with a small lateral denticle at about A collum length

in front of caudal corner; the latter very broadly rounded anteriorly, declined ventrad, caudal corner not

surpassing rear tergal margin.

Body. Tegument smooth and shining, prozonae and surface below paraterga largely finely shagreened,

metaterga rather smooth and leathery, posterior halves faintly rugulose, surface below paraterga finely

microgranulate (Fig. 1A-G). Postcollum metaterga with four transverse rows of very small setigerous

tubercles or setae: an anterior transverse row of 4-6+4-6, intermediate transverse rows (one pre-sulcus.

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European Journal of Taxonomy 195: 1—47 (2016)

the other post-sulcus) of 3—4+3—4, and a posterior transverse row of 5-7+5-7; tergal setae long, strong,

slender, about % of metatergal length (Fig. 1A, D, F). Axial line visible both on pro- and metazonae.

Paraterga well-developed (Fig. 1 A, D, F), especially so in S, lying rather high (at upper l A of body height),

anterior edge rounded and narrowly bordered, fused to callus; caudal corner very narrowly rounded to

pointed. Paraterga 2 and 3 each with two evident setigerous incisions at lateral edge (Fig. 1 A). Following

poreless segments with two evident setigerous incisions; following pore-boring segments often with one

incision in front of pore (Fig. 1C-D). Calluses on paraterga delimited by a sulcus both dorsally and

ventrally. Ozopores evident, lateral, lying in an ovoid groove at about A metatergal length in front of

posterior edge of metaterga. Transverse sulcus usually distinct (Fig. 1A, D, F), slightly incomplete on

segments 4 and 19, complete on segments 5-18, rather wide, line-shaped, rather deep, not reaching

bases of paraterga, beaded at bottom. Stricture between pro- and metazonae evident, wide and rather

deep, ribbed at bottom down to base of paraterga (Fig. 1A, C-F). Pleurosternal carinae complete crests

with a sharp caudal tooth on segments 2 and 3, thereafter increasingly reduced and broken, remaining as

a front bulge and a caudal tooth until segment 10 (c?) or 6(5), on following segments retained as a small

caudal tooth until segment 18 (c?) or absent starting with segment 7 ($). Epiproct (Fig. 1E-G) conical,

flattened dorsoventrally, with two evident, small, rounded apical papillae; tip subtruncate; pre-apical

Fig. 1 . Tylopus flavolineatus sp. nov., holotype, $ (A, C-I), paratype, $ (B). A-B. Anterior part of body,

dorsal and lateral views, respectively. C-D. Segments 10 and 11, dorsal and lateral views, respectively.

E-G. Posterior part of body, lateral, dorsal and subventral views, respectively. H-I. Sternal cones

between coxae 4, caudal and sublateral views, respectively.

4

LIKHITRAKARN N. et al ., The millipede genus Tylopus

lateral papillae evident, lying close to tip. Hypoproct roundly subtriangular, setigerous knobs at caudal

edge small and well-separated (Fig. 1G).

Sterna. Densely setose, with a small, but evident cone caudally near each coxa, rear cones being a

little larger than fore ones (Fig. 2E); an entire, large, tongue-shaped sternal lobe between $ coxae 4

(Fig. 1H, I). Legs rather long and slender, midbody ones ca 1.2-1.3 (c?) or 0.9-1.0 ($) as long as body

Fig. 2. Tylopusflavolineatus sp. nov., holotype, S, left gonopod. A-D. Mesal, lateral, caudal and suboral

views, respectively. E. Sterna of segment 10. F. Leg 6. G. Leg 10. Scale bars = 0.5 mm. Abbreviations

are explained in the text.

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European Journal of Taxonomy 195: 1-47 (2016)

height (Fig. 2G); S femora 5 (Fig. 2F) each with a very strong, distovental, densely pilose adenostyle

in distal l A, S prefemora 10-12 each with a small parabasal adenostyle (Fig. 2G); S prefemora 2 until

segment 12 each with microgranulations; tarsal brushes present until $ leg 6.

Gonopod. Rather simple (Figs 2A-D, 3); coxa slightly curved caudad, sparsely setose distoventrally.

Prefemur as usual, densely setose, about A as long as femorite + postfemoral part. Femorite slightly

curved and rather slender, with an evident mesal groove and a strong distolateral sulcus demarcating

a postfemoral part; lobe 1 rather small, rounded apically, process h large, slightly curved and pointed;

solenophore (sph) clearly curved mesad, long, vaguely bifid, lamina medialis supporting a long

flagelliform solenomere (si).

Remark

In the normal condition, the solenomere is sheathed by the solenophore (Fig. 2A-D), broken off in

Fig. 3.

Fig. 3. Tylopus Jiavolineatus sp. nov., holotype, right gonopod. A-B. Mesal and lateral views,

respectively. C-F. Distal part, subcaudal, subcaudal, suboral and sublateral views, respectively. Scale

bars = 0.2 mm

6

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Tylopus hongkhraiensis sp. nov.

urn:lsid:zoobank.org:act:E217CFlE-83D8-4EFD-875B-7C3845D01580

Figs 4-6, 26

Diagnosis

Differs from all known congeners in the almost missing paraterga, coupled with the gonopod being

especially similar to that of T. magicus Golovatch, 1984, but differs in process h being strong and

Fig. 4. Tylopus hongkhraiensis sp. nov., holotype, S (B-J), paratypes, $ (A). A. Habitus, live

coloration. B-C. Anterior part of body, dorsal and lateral views, respectively. D-E. Segments 10

and 11, dorsal and lateral views, respectively. F-H. Posterior part of body, dorsal, ventral and lateral

views, respectively I-J. Sternal cones between coxae 4, subcaudal and sublateral views, respectively.

Abbreviations are explained in the text.

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European Journal of Taxonomy 195: 1-47 (2016)

pointed, while process z long, slender and pointed.

Etymology

The species is named to emphasize the type locality; adjective.

Material examined

Holotype

THAILAND: S, Huai Hong Khrai Royal Development Study Centre, Doi Salcet, Chiang Mai,

18°52'45" N, 99°12'40" E, ca 420 m a.s.l., 29 Jul. 2015, leg. N. Likhitrakam (CUMZ).

Paratypes

THAILAND: 4 $$, 4 (CUMZ); 1 1 ? (ZMUM p3060); 1 S, 1 $ (ZMUC); 1 S, 1 ? (NHMW),

all same data as for holotype.

Description

Measurements and colour. Length 11.2-12.8 (f) or 11.5-12.6 mm (§), width of midbody pro- and

metazonae 0.85-0.96 and 1.04-1.2 mm (c?) or 0.85-1.05 and 1.05-1.32 mm (5), respectively. Coloration

of live animals dark brown (Pig. 4A); legs light brown, venter and a few basal podomeres light brown to

yellow-brown. Coloration in alcohol after three months of preservation faded to light brown; antennae

and epiproct light brown to pallid, venter and a few basal podomeres light brown to pallid (Pig. 4B-J).

Head. Clypeolabral region and vertex sparsely setose, epicranial suture distinct. Antennae rather short

(Pig. 4A), surpassing body segment 2 (f, 5) when stretched dorsally. In width, segment 3 < 2 < collum

< segment 4<5<6<7< head < 8-17 (f) or segment 2 < 3 < collum < segment 4<5 = 6<7< head <

Fig. 5. Tylopus hongkhraiensis sp. nov., holotype, right gonopod. A-C. Lateral, mesal and suboral

views, respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.

8

LIKHITRAKARN N. et al ., The millipede genus Tylopus

8-17 (5); thereafter body gently and gradually tapering. Collum with three transverse rows of setae:

3+3 anterior, 2+2 intermediate and 3+3 posterior; caudal comer very narrowly rounded, not surpassing

rear tergal margin (Fig. 4B-C).

Body. Tegument smooth and shining, prozonae very finely shagreened, metaterga nearly smooth, faintly

rugulose and leathery (Fig. 4B-F, H). Postcollum metaterga with two transverse rows of setae: 2+2

anterior and 2+2 posterior, traceable at least as insertion points when setae broken off. Tergal setae simple,

slender, rather short, about A of metatergal length. Axial line visible, traceable only on metazonae.

Postcollum paraterga virtually missing, in pore-bearing segments lateral bulges with ozopores set at

about half of midbody height. Ozopores (Op) evident (Fig. 4E), lateral, located at about A of metatergal

Fig. 6. Tylopus hongkhraiensis sp. nov., holotype, left gonopod. A-B. Mesal and lateral views,

respectively. C-F. Distal part, sublateral, submesal, suboral and subcaudal views, respectively. Scale

bars = 0.1 mm.

9

European Journal of Taxonomy 195: 1 —47 (2016)

length in front of posterior edge ofmetaterga. Transverse sulcus complete on segments 6-16, incomplete

on segments 7 and 18 (S, $), very narrow, shallow, not reaching bases of paraterga (Fig. 4B-F, H).

Stricture between pro- and metazonae wide, deep, evidently ribbed at bottom down to base of paraterga

(Fig. 4B-F, H). Pleurosternal carinae complete crests with a very sharp median tooth on segment 2,

increasingly reduced to a rounded caudal crest bulged on segments 2 and 3 {<$, §), retained only as an

evident, sharp, caudal tooth on segments 5 and 6, thereafter increasingly reduced in size until segment

17, remaining as a small caudal tooth on segment 18 (S) (Fig. 4C, E, H), or increasingly reduced to

caudally roughly granulated bulges on segments 5 and 6, thereafter missing (§). Epiproct (Fig. 4F-H)

conical, flattened dorsoventrally, with two small, but evident, rounded apical papillae; tip subtruncate;

lateral pre-apical papillae very small, lying close to tip. Hypoproct roundly subtrapeziform, setigerous

knobs at caudal edge small and well-separated (Fig. 4G).

Sterna. Sparsely setose, without modifications; a small, rounded, densely setose sternal cone between

S coxae 4 (Fig. 41, J). A paramedian pair of small tubercles in front of gonopod aperture. Legs rather

rather short, slightly incrassate in S, midbody ones ca 1.1-1.3 (S) or 0.8-0.9 times ($) as long as body

height, prefemora without modifications, S tarsal brushes absent.

Gonopod. Simple (Figs 5, 6); fermorite faintly curved, stout, slightly expanded distad, showing a strong

mesal groove; process h prominent, slightly curved, pointed; process z long and rather slender, evident,

likewise curved and pointed; solenophore (sph) expanded distally and flattened, typically coiled, clearly

bifid, with a pair of small spines.

Remarks

The millipedes were found mating on bamboo logs on the soil surface during rain. The male was

observed walking along the female’s back and stimulating her with rhythmic pulses of his legs. The tips

of the solenophore and solenomere in Fig. 6 are broken off.

Tylopus moniliformis sp. nov.

um:lsid:zoobank.org:act:A510843C-08CC-4818-A45F-E35AFDE80E71

Figs 7-9, 26

Diagnosis

Differs from all known congeners by the almost missing paraterga, much like in T. hongkhraiensis sp.

nov., coupled with the gonopod structure being similar to that of T. reductus Golovatch, 2013, but it

differs in the solenophore being more strongly twisted and thicker.

Etymology

To emphasize the body being strongly bead-shaped, or moniliform; adjective.

Material examined

Holotype

LAOS: <$, Tad Fane Waterfall, Paksong, Champasak, 15°10'50" N, 106°08'20" E, ca 970 m a.s.l., 20 Jul.

2013, leg. S. Panha, C. Sutcharit, W. Siriwut (CUMZ).

Paratypes

LAOS: 2 33, 8 $$ (CUMZ); 1 <$, 1 $ (ZMUM p3061); 1 & 1 ? (ZMUC); 1 1 $ (NHMW), all

same data as for holotype.

10

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Description

Measurements and colour. Length 14-16 (S) or 16-18 mm (§), width of midbody pro- and metazonae

0.6-0.9 and 0.9-1.1 mm (c?) or 0.9-1.2 and 1.2-1.3 mm (§), respectively. Coloration of live animals

dark castaneous brown (Fig. 7A); legs light brown, venter and a few basal podomeres light brown to

yellow-brown. Coloration of alcohol material after three months of preservation faded to light brown;

antennae and epiproct light brown to pallid, venter and a few basal podomeres light brown to pallid

(Fig. 7B-J).

Fig. 7. Tylopus moniliformis sp. nov., holotype, S (B-J), paratypes, $ (A). A. Habitus, live

coloration. B-C. Anterior part of body, dorsal and lateral views, respectively. D-E. Segments 10 and

11, dorsal and lateral views, respectively. F-H. Posterior part of body, subdorsal, ventral and sublateral

views, respectively. I-J. Sternal cones between coxae 4, subcaudal and sublateral views, respectively.

Abbreviations are explained in the text.

11

European Journal of Taxonomy 195: 1-47 (2016)

Head. Clypeolabral region and vertex sparsely setose, epicranial suture distinct. Antennae moderately

long (Fig. 7A), surpassing body segment 5 (c?) or reaching body segment 4(5) when stretched dorsally.

In width, head < segment 4 < 2 = 3 < collum < segment 5 < 6-17 (f, 5); thereafter body gently and

gradually tapering. Collum with three transverse rows of setae: 5+5 anterior, 2+2 intermediate and 1+1

posterior; a very faint marginal incision laterally in posterior A; caudal corner very narrowly rounded,

not surpassing rear tergal margin (Fig. 7B-C).

Body. Tegument smooth and shining, prozonae very finely shagreened, metaterga almost smooth,

delicately rugulose, leathery (Fig. 7B-F, H). Postcollum metaterga with two transverse rows of setae:

2+2 anterior, always abraded, and 1+1 posterior row, setae traceable at least as insertion points. Tergal

setae simple, strong, slender, about V 3 of metatergal length. Axial line visible, but barely traceable on

prozonae (c?). Paraterga 2 well-developed (Fig. 7B, C), horizontal, anterior edges protruded anteriorly,

fore corner bent ventrad, pointed; lateral edge with two minute incisions in anterior half; caudal corner

very narrowly rounded. Paraterga 3 and 4 rounded, expanded laterally, with two minute incisions only in

segment 3. Following paraterga virtually missing (f) or traceable as small, rounded, laterally expanded

bulges (5), in pore-bearing segments with ozopores set at about half of midbody height. Ozopores

(Op) evident (Fig. 7E), lateral, lying in an ovoid groove at about A of metatergal length in front of

posterior edge of metaterga. Transverse sulcus usually distinct (Fig. 7B-F, H), slightly incomplete on

segment 3, complete on segments 4-18 (c?) or complete on segments 5-18 (5), always incomplete on

segment 19 (<$, 5), narrow, not reaching bases of paraterga, at most faintly beaded at bottom. Stricture

between pro- and metazonae wide, evidently ribbed at bottom down to base of paraterga (Fig. 7B-

Fig. 8. Tylopus moniliformis sp. nov., paratype, left gonopod. A-B. Mesal and lateral views,

respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.

12

LIKHITRAKARN N. et al ., The millipede genus Tylopiis

F, H). Pleurosternal carinae complete crests with a sharp caudal tooth on segments 2 and 3, reduced and

remaining a sharp caudal tooth on segment 4, thereafter missing (S) (Fig. 7C), or thereafter increasingly

reduced and remaining a front bulge until segment 18 (§). Epiproct (Fig. 7F-H) rather short, flattened

dorsoventrally, tip subtruncate, subapical lateral papillae small, but visible, lying close to tip. Hypoproct

roundly subtriangular, setigerous knobs at caudal edge well-separated and evident.

Sterna. Moderately setose, without modifications; an entire, high, inverted funnel-shaped, sternal

lobe each between S coxae 3 and 4 (Fig. 7I-J), lobe being larger between coxae 4. A paramedian

pair of evident tubercles in front of gonopod aperture. Legs very long and slender, slightly incrassate

Fig. 9. Tylopus moniliformis sp. nov., holotype, left gonopod. A-B. Lateral and mesal views,

respectively. C-F. Distal part, lateral, mesal, subcaudal and suboral views, respectively. Scale bars =

0.1 mm.

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European Journal of Taxonomy 195: 1-47 (2016)

in c?, midbody ones ca 1.5-1.8 (c?) or 1.1-1.4 times (§) as long as body height, prefemora without

modifications, S tarsal brushes absent.

Gonopod. Very simple (Figs 8, 9); fermorite straight and rather stout, slightly expanded distad, showing

a distinct mesal groove, without outgrowths except for a low, sometimes poorly delimited lobe 1;

solenophore (sph) expanded and flattened, typically coiled, suberect distally, devoid of processes.

Remarks

This species shows the gonopod with lobe 1 separated from the femorite by a rather indistinct sulcus,

the presence of the latter lobe being one of the main characteristics of Tylopus. Since this locality also

yielded the holotype of Desmoxytes rhinoceros Likhitrakam, Golovatch & Panha, 2014 (cf. Likhitrakam

et al. 2014b), Tad Fane Waterfall becomes a site quite important for nature conservation, being the type

locality of these two millipede species.

Tylopus retusus sp. nov.

urn:lsid:zoobank.org:act:40D74A7C-61D7-442B-B388-3868C47102C5

Figs 10-11,26

Diagnosis

Distinguished by its relatively small body with four transverse rows of setae forming no regular pattern

of setation; in gonopod structure it is similar to T. haplorugosus Golovatch & Enghoff, 1993, but differs

in process h being rather short, slightly flattened, blunt, coupled with the solenophore being strongly

coiled.

Etymology

To emphasize the blunt tip of process h; adjective.

Material examined

Holotype

LAOS: Muang Xay City, Xay, Oudomxai, 20°41'00" N, 101°59'05" E, ca 640 m a.s.l., mango garden,

14 Oct. 2014, leg. C. Sutcharit & R. Srisonchai (CUMZ).

Paratype

LAOS: 1 5, Khoua, Phongsaly, ca 835 m a.s.l., 21°H'55" N, 102°06'40" E, forest near road, 15 Oct.

2014, leg. C. Sutcharit & R. Srisonchai (CUMZ).

Description

Measurements and colour. Length 16.3 (S) or 20.4 mm (§), width of midbody pro- and metazonae 1.12

and 2.07 mm (f) or 2.7 and 3.7 mm (§), respectively. Live coloration light brown (Fig. 10A); prozonae

dark brown; paraterga, antennae, head and legs light brown to yellow-brown. Coloration of alcohol

material after one year of preservation faded to light brown to yellow-brown; paraterga, antennae, head,

legs and venter light brown to light yellow (Fig. 10B-J).

Head. Clypeolabral region and vertex sparsely setose, epicranial suture distinct. Antennae (Fig. 10A,

B) rather short and stout, antennomeres 2-6 subequal in length, reaching body segment 3 (f, $) when

stretched dorsally. In width, head < segment 3 < 4 < collum < segments 5-15; thereafter body gently

and gradually tapering. Collum with three transverse rows of setae: 6+6 anterior, 4+4 intermediate and

6+6 posterior; a small lateral incision at about midway; caudal comer very broadly rounded, paraterga

declined ventrad, slightly produced behind rear tergal margin.

14

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Body. Tegument smooth and shining, prozonae finely shagreened, metaterga leathery, finely rugulose

(Fig. 10B, D, F); surface below paraterga roughly microgranulate (Fig. IOC, E, H). Postcollum

metaterga with four transverse rows of setae forming no regulated pattern of setation, anterior row with

the most dense setae growing increasingly dense towards segment 19 (Fig. 10B, D, F). Tergal setae

long and slender, about % as long as metaterga. Axial line well visible on metazonae, traceable also on

prozonae. Paraterga strongly developed (Fig. 10B-H), especially so in S, set rather high (at upper V 3 of

body height), slightly upturned, but lying below dorsum; anterior edge broadly rounded and narrowly

bordered, fused to callus; caudal corner very narrowly rounded, starting with segment 16 extending

Fig. 10. Tylopus retusus sp. nov., holotype, $. A. Habitus, live coloration. B-C. Anterior part of body,

dorsal and lateral views, respectively. D-E. Segments 10 and 11, dorsal and lateral views, respectively.

F-H. Posterior part of body, dorsal, ventral and lateral views, respectively. I-J. Sternal cones between

coxae 4, caudal and sublateral views, respectively.

15

European Journal of Taxonomy 195: 1-47 (2016)

increasingly behind rear tergal margin (Fig. 10F, H). Paraterga 2 and 3 with two evident incisions at

lateral edge (Fig. 10B). Following poreless segments with two similar incisions; pore-boring segments

with one, often setigerous incision lying in front of pore (Fig. 10D, E). Calluses of paraterga delimited

Fig. 11. Tylopus retusus sp. nov., holotype, $, left gonopod. A-D. Mesal, lateral, subcaudal and suboral

views, respectively. E. Sterna of segment 10. F. Leg 6. G. Leg 10. Scale bars = 0.2 mm. Abbreviations

are explained in the text.

16

LIKHITRAKARN N. et al ., The millipede genus Tylopus

by a sulcus both dorsally and ventrally. Ozopores evident, lateral, lying in an ovoid groove at about A of

metatergal length in front of posterior edge of metaterga. Transverse sulcus usually distinct (Fig. 10B,

D, F), slightly incomplete on segment 19, complete on segments 5-18, narrow, undulate, shallow, not

reaching bases of paraterga, beaded at bottom. Stricture between pro- and metazonae evident, wide

and deep, beaded at bottom down to base of paraterga (Fig. 10B, D-F). Pleurosternal carinae complete

crests with a sharp caudal tooth on segments 2-7 (c?) or 2-4 (?), thereafter increasingly reduced to

a small tooth until segment 15 (c?) or 13 (?), missing more caudally. Epiproct (Fig. 10F-H) conical,

flattened dorsoventrally, with two evident, large, rounded apical papillae; tip subtruncate; lateral pre-

apical papillae small, lying close to tip. Hypoproct (Fig. 10G) roundly subtrapeziform, setigerous knobs

at caudal edge small and well-separated.

Sterna. Densely setose, with a small cone caudally near each coxa, rear cones being a little larger than

fore ones (Fig. 11E); two small, rough, fully separated, setose cones between $ coxae 4 (Fig. 101, J).

Legs rather long and slender, midbody ones ca 1.2-1.3 {$) or 0.9-1.0 (?) as long as body height;

all male legs until segment 16 with tarsal brushes, prefemora not swollen; prefemora, femora and

tibiae with ventral microgranulations; femora, tibiae and tarsi each with an evident adenostyle (tubercle)

medially (Fig. 11F-G).

Gonopod. Rather simple (Fig. 11A-D); coxa a little curved caudad, sparsely setose distoventrally.

Femorite slightly curved and stout, with an evident mesal groove and a clear distolateral sulcus

demarcating a postfemoral part; lobe 1 evident, rounded, with process h rather short, slightly flattened,

blunt; solenophore (sph) strongly coiled, devoid of processes, pointed.

Remark

This species was found in a mango plantation in Muang Xay City.

Tylopus acuminatus sp. nov.

urn:lsid:zoobanlc.org:act:C0DB482F-EE58-4385-9733-7D2E724DEFED

Figs 12-15, 26

Diagnosis

Very similar to T. jeekeli Golovatch & Enghoff, 1993 and T. parajeekeli Likhitrakarn, Golovatch,

Prateepasen & Panha, 2010, especially in its gonopod conformation, but differs in process h being

higher and more strongly flattened, pointed, while the sternal lobe between S coxae 4 linguiform and

larger.

Etymology

To emphasize the acuminate processes h on the gonopod; adjective.

Material examined

Holotype

LAOS: S, Ban Sin Chai, Boun Neua, Phongsali, 21°31'50" N, 101°51'45" E, ca 820 m a.s.l., 15 Oct.

2014, leg. C. Sutcharit & R. Srisonchai (CUMZ).

Paratypes

LAOS: 1 S, 1 ?, Phuthalang forest protected area, Boun Neua, Phongsali, 21°37'30" N, 101°55'55"

E, ca 1125 m a.s.l., 16 Oct. 2014 (CUMZ); 1 c?, 2 ??, Phufa, Boun Neua, Phongsali, 21°41'05" N,

102°06'35" E, ca 1470 m a.s.l, 16 Oct. 2014 (CUMZ); 1 ?, near Ban Borkhun, Boun Neua, Phongsali,

21°27'57" N, 101°47'20" E, ca 880 m a.s.l., 18 Oct. 2014 (CUMZ); 1 ?, Khoua, Phongsali, 21° 1F55'" N,

102°06'40" E, ca 840 m a.s.l., 15 Oct. 2014 (CUMZ); 1 S, 1 ? (NHMW), 1 S (ZMUM p3059), BanNa

17

European Journal of Taxonomy 195: 1-47 (2016)

Thong, Namo, Oudomxay, 20°52'25" N, 101°47'00" E, ca 650 m a.s.l., 14 Oct. 2014; 1 S, Phagneung

Phoukulom Waterfall, Sing, Luang Namtha, 21°07T5" N, 101°14'40" E, ca 900 m a.s.l., 13 Oct. 2014,

all leg. C. Sutcharit & R. Srisonchai (ZMUC).

Description

Measurements and colour. Length 24.5-31.6 (c?) or 24.3-31.5 mm (J), width of midbody pro- and

metazonae 1.98-2.6 and 2.95-3.6 mm (c?) or 2.7-2.9 and 3.1—4.2 mm (§), respectively. Live coloration

dark brown (Pig. 12A); antennae and epiproct light brown; legs yellow-brown. Coloration of alcohol

Fig. 12. Tylopus acuminatus sp. nov., holotype, S- A. Habitus, live coloration. B-C. Anterior part

of body, dorsal and lateral views, respectively. D-E. Segments 10 and 11, dorsal and lateral views,

respectively. F-H. Posterior part of body, dorsal, ventral and lateral views, respectively. I-J. Sternal

cones between coxae 4, caudal and sublateral views, respectively.

18

LIKHITRAKARN N. et al ., The millipede genus Tylopus

material after one year of preservation faded to brown to yellow-brown; head, antennae, epiproct light

brown to yellow-brown, legs and venter yellow-brown to light yellow (Fig. 12B-J).

Head. Clypeolabral region densely, vertex sparsely, setose; epicranial suture distinct. Antennae

(Fig. 12A) short, surpassing body segment 2 (<$, $) when stretched dorsally. In width, head < segment

3 < collum < segment 4 < 2 < 5-16 (c?), or head < segment 3 < 4 < collum < segment 2 < 5-16 (?);

thereafter body gently and gradually tapering. Collum with three transverse rows of setae: 4+4 anterior,

1+1 intermediate and 2+2 posterior paraterga declined ventrad, with a small lateral incision at about

midway; caudal comer very broadly rounded, not surpassing rear tergal margin (Fig. 12C).

Body. Tegument smooth and shining, prozonae finely shagreened, metaterga smooth and leathery

(Fig. 12B, D, F); surface below paraterga microgranulate (Fig. 12C, E, H). Postcollum metaterga with

an anterior transverse row of 2+2, mostly abraded setae; posterior row barely traceable as 4+4 insertion

points, these being better visible on segments 17-19 (Fig. 12F). Tergal setae simple and slender, about

% as long as metaterga. Axial line traceable on prozonae and anterior halves of metazonae. Paraterga

well-developed (Fig. 12B-F, H), especially so in S, set rather high (at upper A of body height), slightly

upturned, but lying below dorsum; anterior edge broadly rounded and narrowly bordered, fused to

callus; caudal corner very narrowly rounded, drawn behind tergal margin, increasingly protruding and

pointed starting with segment 16 (Fig. 12F, H). Paraterga 2 with two evident incisions at lateral edge

in anterior A. Paraterga 3 and 4 each with two similar incisions at lateral edge, one in anterior A, the

other about the middle. Following paraterga each with one evident lateral incision in anterior A and,

in pore-bearing segments, a very small denticle in the middle (Fig. 12B, D, F). Calluses on paraterga

delimited by a sulcus both dorsally and ventrally. Ozopores evident, lateral, lying in an ovoid groove

Fig. 13. Tylopus acuminatus sp. nov., holotype, right gonopod. A-B. Tateral and mesal views,

respectively. C. Teg 13. Scale bar: A-B. 0.5 mm; C = no scale bar. Abbreviations are explained in the

text.

19

European Journal of Taxonomy 195: 1—47 (2016)

at about l A of metatergal length in front of posterior edge of metaterga. Transverse sulcus evident on

metaterga 5-17, incomplete on metaterga 18, narrow, line-shaped, rather deep, ribbed at bottom, not

reaching bases of paraterga (Fig. 12B, D, F). Stricture between pro- and metazonae evident, wide and

deep, ribbed at bottom down to base of paraterga (Fig. 12B, D-F). Pleurosternal carinae complete crests

with a sharp caudal tooth on segments 2 and 3 (<$, $), increasingly reduced and remaining a strong,

sharp, caudal tooth on segments 4-8, retained as a small sharp tooth on segments 9-16, a minute tooth

on segment 17 (c?), or an evident, sharp, caudal tooth on segments 4-15, then remaining a small rounded

tooth on segment 16, thereafter missing (§) (Fig. 12C, E, H). Epiproct (Fig. 12F-H) conical, flattened

dorsoventrally, with two evident, rounded, apical papillae; tip subtruncate; lateral pre-apical papillae

Fig. 14. Tylopus acuminatus sp. nov., holotype, right gonopod. A-B. Mesal and lateral views,

respectively. C-F. Distal part, sublateral, submesal, subcaudal and suboral views, respectively. Scale

bars = 0.2 mm

20

LIKHITRAKARN N. et al ., The millipede genus Tylopus

evident, lying close to tip. Hypoproct (Fig. 12G) roundly subtrapeziform, setigerous knobs at caudal

edge small and well-separated.

Sterna. Moderately setose and densely microtuberculate; cross-impressing shallow; an entire,

linguiform, sternal lobe between $ coxae 4 (Fig. 121—J). A paramedian pair of conspicuous ridges in

front of gonopod aperture. Legs rather long and slender, midbody ones ca 1.2-1.3 (c?) or 1.0-1.1 (§)

as long as body height; $ legs with prefemora distinctly swollen laterally, acropodites with particularly

dense setae ventrally (Fig. 13C), but tarsal brushes absent.

Fig. 15. Tylopus acuminatus sp. nov., paratype, from Phufa (A-B), paratype, c?, from Phuthalang

forest protected area (C-D), paratype, from forest near road (E), paratype, from Ban Na Thong

(F), paratype, from Phagneung Phoukulom waterfall (G-H). A-H. Distal part, sublateral, submesal,

dorsal, subdorsal, dorsal, subdorsal, ventral and dorsal views, respectively. Scale bars = 0.2 mm. Red

colour shows process h. Green colour shows spine z.

21

European Journal of Taxonomy 195: 1—47 (2016)

Gonopod. Rather simple (Figs 13A-B, 14-15); coxa slightly curved caudad, sparsely setose

distoventrally. Femorite slightly curved and stout, with an evident mesal groove and a clear distolateral

sulcus demarcating a postfemoral part; process h prominent, evidently curved, pointed, with a small,

but evident dorsal tooth and sometimes with a small ventral denticle (Fig. 13A-B); spine z pointed,

sometimes small (Fig. 15G); solenophore (sph) large, expanded, flattened and typically coiled.

Remark

This species shows slight variations in gonopod structure, process h often being rather narrow to slightly

broader, while spine z small to slightly broader (Fig. 15).

Tylopus dorsalis sp. nov.

urn:lsid:zoobank.org:act:EDC8D353-BE43-4D79-8BA4-E23D0FD57CE7

Figs 16-17, 26

Diagnosis

The live coloration is similar to that of T. trigonum , but the new species differs in the head and collum

being light brown (versus blackish), and the pattern not contrasting (versus contrasting); the gonopod

structure is similar to that of I crassipes Golovatch, 1984, but the new species differs in process x being

longer and its tip rounded dorsally.

Etymology

To emphasize process x on the solenophore arising dorsally; adjective.

Material examined

Holotype

LAOS: S, Phuthalang forest protected area, Boun Neua, Phongsali, 21°37'30" N, 101°55'55" E, ca

1125 m a.s.l., 16 Oct. 2014, leg. C. Sutcharit & R. Srisonchai (CUMZ).

Paratype

LAOS: 1 5, same data as for holotype (CUMZ).

Description

Measurements and colour. Length 22.1 ( f ) or 22.3 mm (§), width of midbody pro- and metazonae

2.5 and 3.6 mm (c?) or 1.97 and 2.55 mm (§), respectively. Live coloration light brown (Fig. 16A);

paraterga, legs and epiproct light yellow-brown, head and collum light brown, antennomere 7 blackish,

collum and following terga each with a light brown triangle and a dark brown collar covering both pro-

and metazonae. Coloration of alcohol material faded after one year of preservation to light brown, with

a pattern of a dark brown band in pre-sulcus area on metazonae and a pair of paramedian, longitudinal,

brown stripes dorsally both on pro- and metaterga; paraterga light red-brown, head brown, antennae,

legs, venter and epiproct light yellow-brown (Fig. 16B-J).

Head. Clypeolabral region densely, vertex sparsely, setose; epicranial suture distinct. Antennae rather

short (Fig. 16A), extending behind body segment 3 (S) or 2 ($) when stretched dorsally. In width, head

< segment 3 < 2 = 4 < collum < segments 5-17 (S, $); thereafter body gently and gradually tapering.

Collum with three transverse rows of setae: 3+3 anterior, 1+1 intermediate and 2+2 posterior; a small

lateral denticle at about anterior A (Fig. 16B, D); caudal corner of paraterga very broadly rounded,

declined ventrad, not surpassing rear tergal margin.

22

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Body. Tegument smooth and shining, prozonae finely shagreened, metaterga smooth and leathery,

posterior halves rugulose, surface below paratergamicrogranulate (Fig. 16B-F, H). Postcollum metaterga

with two transverse rows of setae: 2+2 in anterior (pre-sulcus) row, also 2+2 in posterior (post-sulcus)

one. Tergal setae long, strong, slender, about % of metatergal length. Axial line traceable both on pro- and

metazonae. Paraterga well-developed (Fig. 16B, D, F), especially so in S, set rather high (at upper l A of

body height), anterior edge rounded and narrowly bordered, fused to callus; caudal corner very narrowly

rounded, extending increasingly behind rear tergal margin, posterior edge slightly oblique. Calluses on

paraterga delimited by a sulcus only dorsally. Paratega 2 broad, anterior edge angular, lateral edge with

Fig. 16. Tylopus dorsalis sp. nov., holotype, $. A. Habitus, live coloration. B-C. Anterior part of body,

dorsal and lateral views, respectively. D-E. Segments 10 and 11, dorsal and lateral views, respectively.

F-H. Posterior part of body, dorsal, ventral and lateral views, respectively. I-J. Sternal cones between

coxae 4, caudal and sublateral views, respectively.

23

European Journal of Taxonomy 195: 1-47 (2016)

three evident incisions. Lateral edge of following paraterga with a clear incision in anterior Vs (Fig. 16B,

D). Paraterga 14-19 with caudal corner extending increasingly behind, with tip evidently curved mesad

(Fig. 16F). Ozopores distinct, lateral, lying in an ovoid groove at about Vs of metatergal length in front

of posterior edge of metaterga. Transverse sulcus usually distinct (Fig. 16B, D, F), slightly incomplete

on segment 3, complete on segments 4-18, narrow, line-shaped, deep, not reaching bases of paraterga,

beaded at bottom. Stricture between pro- and metazona evident, wide, rather deep, clearly ribbed at

bottom down to base of paraterga (Fig. 16B, D-G). Pleurosternal carinae complete crests with a sharp

Fig. 17. Tylopus dorsalis sp. nov., holotype, f. A-D. Right gonopod, lateral, mesal, caudal and oral

views, respectively. Scale bars = 0.2 mm. Abbreviations are explained in the text.

24

LIKHITRAKARN N. et al ., The millipede genus Tylopus

caudal tooth on segments 2-7, reduced and remaining only a sharp caudal tooth on segment 8, reduced

to a small, delicately denticulate, caudal crest on segments 9-15, retained as a minute tooth on segment

16, thereafter missing (c?), or with complete crests and a sharp caudal tooth on segments 2^1, remaining

only an evident caudal tooth on segments 5-12, a rounded caudal crest on segments 13-17, thereafter

missing (§) (Fig. 16C, E, H). Epiproct (Fig. 16E-G) conical, flattened dorsoventrally, with two evident,

rounded, apical papillae; tip subtruncate; lateral pre-apical papillae evident, lying close to tip. Hypoproct

roundly subtriangular, setigerous knobs at caudal edge small and well-separated (Fig. 16G).

Sterna. Separsely setose, without modifications; cross-impressions shallow; an entire, linguiform,

sternal lobe between $ coxae 4 (Fig. 161—J). Legs long and slender, midbody ones ca 1.3-1.4 (c?) or

0.9-1.1 (?) as long as body height; $ femora 6 bulged and microtuberculate ventrally (Fig. 17E); $

femora 7 also bulged ventrally (Fig. 17F), but following femora not swollen; legs of segments 7-16

each with femur, postfemur, tibia and tarsus carrying a small adenostyle (tubercle) medially (Fig. 17G),

telopodites particularly densely setose ventrally (Fig. 17G), tarsal brushes present until $ segment 6.

Gonopod. Rather simple (Fig. 17A-D); coxa slightly curved caudad, sparsely setose distoventrally.

Femorite slightly curved and stout, with an evident mesal groove and a clear distolateral sulcus

demarcating a postfemoral part; process h rather high, directed anterodorsally, flattened, tip rounded;

solenophore (sph) rather slender, strongly coiled, flattened and evidently bifid, with process x being slim

and directed dorsally.

Tylopus thunghaihin sp. nov.

urn:lsid:zoobank.org:act:D8C75297-3063-4B 15-AD A6-D86AD5796B96

Figs 18-20, 26

Diagnosis

The new species seems to be especially similar to T. nodulipes (Attems, 1953), but differs in the body

tegument being dull, coupled with gonopodal process m smaller, process z high and rounded, and

process x pointed.

Etymology

To emphasize the type locality, in Lao “Thung Hai FI in”, commonly translated as the famous Plain of

Jars, a noun in apposition.

Material examined

Holotype

LAOS: S, Plain of Jars, Xieng Khouang, 19°25'50" N, 103°09T5" E, ca 1100 m a.s.l., 26 Nov. 2014,

leg. S.I. Golovatch, C. Sutcharit, N. Likhitralcam (CUMZ).

Paratype

LAOS: 1 5, same data as for holotype (CUMZ).

Description

Measurements and colour. Length 26.1 (c?) or 26.2 mm ($), width of midbody pro- and metazonae

2.02 and 3.02 mm (3) or 2.04 and 3.04 mm (§), respectively. Live coloration brown black (Fig. 18A);

antennae, paraterga and legs dark brown. Coloration in alcohol after one year of preservation blackish,

paraterga dark brown; venter and a few basal podomeres light brown to yellow-brown, legs increasingly

darker brown distally (Fig. 18B-J).

25

European Journal of Taxonomy 195: 1-47 (2016)

Head. Clypeolabral region and vertex sparsely setose, epicranial suture distinct. Antennae rather short

(Fig. 18A), extending behind body segment 3 (c?) or 2 (§) when stretched dorsally. In width, head <

segment 3 < 4 < 5 < collum < segment 2 < 6-16 (<$, $); thereafter body gently and gradually tapering.

Collum with three transverse rows of setae: 4+4 anterior, 1+1 intermediate and 3+3 posterior; with a

small lateral incision near midway (Fig. 18B, D); caudal corner of paraterga very broadly rounded,

declined ventrad, not surpassing rear tergal margin.

Fig. 18. Tylopus thunghaihin sp. nov., holotype, $. A. Habitus, live coloration B-C. Anterior part

of body, dorsal and lateral views, respectively. D-E. Segments 10 and 11, dorsal and lateral views,

respectively. F-H. Posterior part of body, dorsal, ventral and lateral views, respectively. I-J. Sternal

cones between coxae 4, caudal and sublateral views, respectively.

26

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Body. Tegument dull, shining, prozonae finely shagreened; metaterga often roughly microgranulate and

rugulose, leathery; surface below paraterga more delicately, but still sufficiently clearly microgranulate

and rugulose (Fig. 18A-F, H). Metaterga with two transverse rows of small setigerous cones: 2+2 in

anterior (pre-sulcus) row and 3+3 setae usually borne on low, oblong, rounded tubercles in posterior

(post-sulcus) row, except for metaterga 17-19, these latter bearing 3(2)+3(2) setae in anterior row and

4+4 in posterior row. Tergal setae simple, slender, often abraded, about as long as Vs of metazonite. Axial

line visible both on pro- and metazonae. Paraterga strongly developed (Fig. 18B, D, F), especially so in

S, set high (at upper % of body height), anterior edge rounded and narrowly bordered, fused to callus;

caudal corner very narrowly rounded, extending increasingly behind rear tergal margin, posterior edge

slightly oblique. Calluses on paraterga delimited by a sulcus only dorsally. Paratega 2 broad, anterior

edge angular, lateral edge with three evident incisions. Lateral edge of following paraterga with a clear

incision in anterior Vs (Fig. 18B, D, F). Paraterga 16-19 with caudal corner extending increasingly

caudally, tip clearly curved mesad (Fig. 18F). Ozopores evident, lateral, lying in an ovoid groove at

about Vs in front of posterior edge of metaterga. Transverse sulcus distinct (Fig. 18B, D, F), slightly

incomplete on segments 4 and 18, complete on segments 5-17, narrow, wave-shaped, rather deep, not

reaching bases of paraterga, ribbed at bottom. Stricture between pro- and metazonae evident, wide,

deep, ribbed at bottom down to base of paraterga (Fig. 18B-G). Pleurosternal carinae complete crests

with a sharp caudal tooth on segments 2-4 (<$, $), increasingly reduced to a front bulge and a caudal

tooth until segment 7, retained only as a sharp caudal tooth until segment 16, thereafter missing (c?), or

increasingly reduced and remaining only a sharp caudal tooth until segment 15, thereafter missing (§)

(Fig. 18C, E, H). Epiproct (Fig. 18F-H) conical, flattened dorsoventrally, with two evident, rounded.

Fig. 19. Tylopus thunghaihin sp. nov., holotype, $. A-B. Right gonopod, lateral and mesal views,

respectively. C. Leg of segment 9. Scale bars: A-B = 0.2 mm; C = 1 mm. Abbreviations are explained

in the text.

27

European Journal of Taxonomy 195: 1-47 (2016)

apical papillae; tip subtruncate; lateral pre-apical papillae large, lying close to tip. Hypoproct roundly

subtriangular, setigerous knobs at caudal edge small and well-separated (Fig. 18G).

Sterna. Sparsely setose, with a small, but evident cone caudally near each coxa, rear cones being a little

larger than fore ones (Fig. 19C); cross-impressions shallow; a large, central, slightly bifid, setose lobe

between S coxae 4 (Fig. 181—J). Legs long and slender, midbody ones ca 1.2-1.4 (f) or 1.1-1.2 (§) as

long as body height; legs of S segments 5-17 with femur showing a field of ventral microgranulations,

while postfemur, tibia and tarsus each with 1-3 minute adenostyles (tubercles), prefemora not swollen

(Fig. 19G), S telopodites with particularly dense short setae ventrally (Fig. 19G), tarsal brushes present

until body segment 8.

Fig. 20. Tylopus thunghaihin sp. nov., holotype, S, right gonopod. A-B. Mesal and lateral views,

respectively. C-F. Distal part, submesal, sublateral, subcaudal and suboral views, respectively. Scale

bars = 0.2 mm

28

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Gonopod. Rather complex (Figs 19A-B, 20); coxa slightly curved caudad, sparsely setose distoventrally.

Femorite slightly curved and stout, with an evident mesal groove and a clear distolateral sulcus

demarcating a postfemoral part; process h high, strongly twisted, tip evidently bifid; process m short;

process z rather long, sausage-shaped; solenophore (sph) large, strongly coiled, expanded distally,

flattened and clearly bifid; process x large, dentiform and pointed dorsally.

Remarks

This species has been found under stones, each specimen located in its own burrow of ca 30 cm long

under its own stone and forming a kind of small cavern, all in the Plain of Jars, Taos.

Tylopus punctus sp. nov.

urn:lsid:zoobank.org:act:35585ACA-8490-49EE-8DA5-73EB3EB04A5C

Figs 21-22, 26

Diagnosis

Very similar to T. silvestris (Pocock, 1895), but differs in the smaller size, the width of midbody being

2.1-2.5 mm (versus 3.0 mm); the caudal corner of midbody paraterga very narrowly rounded and not

drawn behind the rear tergal margin (versus nearly pointed and produced behind the rear tergal margin),

and gonopod process h small and pointed (versus absent).

Etymology

To emphasize gonopod process h being small and pointed; adjective.

Material examined

Holotype

MYANMAR: S, Mintaingbin Forest Camp, ca 35 km north of Aungban, Chan State, 20°55'20" N,

96°33'60" E, ca 1300 m a.s.l., 11-23 Jun. 2004, leg. H. Schillhammer & H.V. Shaverdo (NHMW-

8248).

Paratype

MYANMAR: 1 §, same data as for holotype (NHMW-8249).

Description

Measurements and colour. Length 27.5 (c?) or 29 mm (§), width of midbody pro- and metazonae 1.56

and 2.1 mm (c?) or 1.98 and 2.5 mm (§), respectively. Coloration of alcohol material upon long-term

preservation rather uniformly light brown to brown, with light yellow paraterga, antennae, venter and

legs (Fig. 21A-I).

Head. Clypeolabral region and vertex sparsely setose, epicranial suture distinct. Antennae rather short

(Fig. 2 IB), reaching only in front of body segment 3 (S, §) when stretched dorsally. In width, segment

3 < 4 < head < collum < segment 2 < 5-15; thereafter body gently and gradually tapering. Collum with

three transverse rows of setae: 4+4 anterior, 2+2 intermediate and 3+3 posterior; an evident lateral

incision at about anterior A; caudal corner of paraterga very broadly rounded, declined ventrad, not

drawn behind rear tergal margin.

Body. Tegument smooth and shining, prozonae very finely shagreened, metaterga rather smooth

and leathery, with posterior halves faintly rugulose, surface below paraterga finely microgranulate

(Fig. 21A-F). Postcollum metaterga with two transverse rows of setae: 2+2 in anterior (pre-sulcus)

row, mostly abraded, and 2+2 in posterior (post-sulcus) row, mostly traceable as insertion points, except

29

European Journal of Taxonomy 195: 1—47 (2016)

for segments 18 and 19, each latter with 4+4 setae in posterior row. Tergal setae simple, slender, about

l A of metatergal length (Fig. 21 A, F). Axial line visible both on pro- and metazonae. Paraterga well-

developed (Fig. 21A-G), especially so in set high (at upper % of body height), anterior edge rounded

and narrowly bordered, fused to callus; caudal corner very narrowly rounded, segments 2 and 3 slightly

extending behind rear tergal margin, thereafter not surpassing rear tergal margin until segment 15

(Fig. 21A-D). Paratega 2 broad, anterior edge angular, lateral edge with two small, but evident incisions

in anterior half. Following paraterga with an evident incision in anterior A of lateral edge (Fig. 21 A,

C, F). Calluses on paraterga delimited by a sulcus only dorsally, rather narrow, a little wider on pore¬

bearing segments. Ozopores distinct, lying at about % of metatergal length in front of posterior edge

of metaterga. Transverse sulcus usually distinct (Fig. 21 A, D, F), slightly incomplete on segment 19,

complete on segments 5-18, narrow, line-shaped, rather deep, reaching bases of paraterga, beaded at

bottom. Stricture between pro- and metazonae evident, rather wide and deep, beaded at bottom down

to base of paraterga (Fig. 21 A, C-F). Pleurostemal carinae complete crests with a sharp caudal tooth

on segments 2 and 3, thereafter increasingly reduced and remaining a sharp caudal tooth until segment

9 (c?) or 6 ($), retained as a small caudal tooth and increasingly reduced until segment 17, absent from

segment 18 (c?) or 7 ($). Epiproct (Fig. 21E-G) conical, flattened dorsoventrally, with two evident.

Fig. 21. Tylopus punctus sp. nov., holotype, f. A-B. Anterior part of body, dorsal and lateral views,

respectively. C-D. Segments 11-12, dorsal and lateral views, respectively. E-G. Posterior part of body,

lateral, subdorsal and subventral views, respectively. H-I. Sternal cones between coxae 4, subcaudal

and sublateral views, respectively.

30

LIKHITRAKARN N. et al ., The millipede genus Tylopus

rather large, rounded, apical papillae; tip subtruncate; lateral pre-apical papillae clear, lying close to tip.

Hypoproct roundly subtriangular, setigerous knobs at caudal edge small and well-separated (Fig. 21G).

Sterna. Sparsely setose, without modifications, but with a pair of small, blunt, fully separated cones

between S coxae 4 (Fig. 21H, I). Legs rather short and slender, midbody ones ca 1.1-1.2 (c?) or 0.8-0.9

( 5 ) as long as body height, $ prefemora not swollen, $ tarsal brushes present only on legs 1 and 2.

Gonopod. Rather simple (Fig. 22); coxa slightly curved caudad, sparsely setose distoventrally. Prefemur

sparsely setose, about A as long as femorite + postfemoral part. Femorite curved and stout, with an

evident mesal groove and a strong distolateral sulcus demarcating a postfemoral part; process h small,

slender and pointed; solenophore (sph) clearly coiled, long and flattened.

Remark

Pocock (1895) described only T. doriae (Pocock, 1895) and T. silvestris from northern Myanmar, the

above being a third species of this genus also occurring in northern Myanmar.

Fig. 22. Tylopuspunctus sp. nov., holotype, <$■ A-D. Left gonopod, lateral, mesal, subcaudal and suboral

views, respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.

31

European Journal of Taxonomy 195: 1-47 (2016)

New faunistic record

Tylopus baenzigeri Golovatch & Enghoff, 1993

Figs 23-26

Tylopus baenzigeri Golovatch & Enghoff, 1993: 101, figs 50-51 (Type locality, Doi Suthep National

Park).

Tylopus baenzigeri -Enghoff 2005: 99. —Likhitrakam et al. 2010: 25; 2014a: 65.

Fig. 23. Tylopus baenzigeri Golovatch & Enghoff, 1993, $. A. Habitus, live coloration. B-C. Anterior

part of body, dorsal and lateral views, respectively. D-E. Segments 10 and 11, dorsal and lateral views,

respectively. F-H. Posterior part of body, dorsal, ventral and lateral views, respectively. I-J. Sternal

cones between coxae 4, subcaudal and sublateral views, respectively.

32

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Material examined

THAILAND: S, Royal Agricultural Station Angkhang, Fang, Chiang Mai, 19°54’05”N, 99° 2’25”E,

ca 1420 m a.s.l., 12 August 2014, leg. N. Likhitrakam (CUMZ).

Descriptive notes

Measurements and colour. Length 43.8 mm, width of midbody pro- and metazonae 3.51 and 5.04 mm,

respectively. Coloration of live animal, as well as of alcohol material black-brown (Fig. 23A-G), head

and venter dark to light brown, legs and antennae yellowish brown to light yellowish.

Body. Antennae rather short (Fig. 23B), extending behind body segment 3 (c?) when stretched dorsally.

In width, segment 3 < 4 < 5 < collum < segment 2 < 6-17; thereafter body gently and gradually tapering.

Collum with three transverse rows of setae: 4+4 anterior, 1+1 intermediate and 4+4 posterior; a very

faint incision laterally in posterior A; caudal corner of paraterga very broadly rounded, not drawn behind

rear tergal margin (Fig. 23B-C). Paraterga 2 well-developed (Fig. 23B-C), horizontal, lateral edge with

two minute incisions in anterior half, caudal corner very narrowly rounded. Paraterga 3 and 4 each

nearly pointed, expanded laterally, with two minute incisions in the middle. Sterna densely setose and

finely microtuberculate (Fig. 24D); with a large, subtrapeziform, ventrally slightly concave, sternal lobe

between <$ coxae 4 (Fig. 23I-J). Legs rather long and slender, midbody ones ca 1.2-1.3 (c?) as long as

Fig. 24. Tylopus baenzigeri Golovatch & Enghoff, 1993, S- A-B. Right gonopod, lateral and mesal

views, respectively. C. Leg 8. D. Legs of segment 11. Scale bars = 0.5 mm. Abbreviations are explained

in the text.

33

European Journal of Taxonomy 195: 1—47 (2016)

body height; 3 legs without tarsal brushes; 3 postfemora and tibiae each with an adenostyle (tubercle)

on segments 8-17; $ coxae, prefemora, femora and postfemora microtuberculate ventrally (Fig. 24C-D).

Remarks

This new specimen agrees in most details with the original description (Golovatch & Enghoff 1993),

especially in gonopod structure which shows a huge spine z serrate on both sides (Figs 24A-B, 3),

and in a large body. Thus, to document its identity the new specimen has been illustrated in detail for

comparative purposes (Figs 23-25). The type locality of T. baenzigeri , Doi Suthep National Park, is

located quite far away (about 120 km) from this new place.

Checklist

Table 1 presents a checklist of the known or new Tylopus species, all arranged in alphabetic order. Data

on their distribution are also provided.

Fig. 25. Tylopus baenzigeri Golovatch & Enghoff, 1993, 3, right gonopod. A-B. Mesal and lateral

views, respectively. C-F. Distal part, submesal, sublateral, subcaudal and suboral views, respectively.

Scale bar = 0.2 mm.

34

LIKHITRAKARN N. et al ., The millipede genus Tylopus

Table 1. Records of Tylopus species.

No.

Species

Localities

1

Tylopus acuminatus sp. nov.

Ban Sin Chai; Phuthalang forest protected

area; Phufa; Ban Borkhun; Khoua; Ban Na

Thong, Phongsali Province, Phagneung

Phoukulom waterfall, Luang Namtha

Province, Laos (see details above).

2

Tylopus affinis Golovatch & Enghoff, 1993

Doi Suthep National Park (Golovatch &

Enghoff 1993); Doi Inthanon National Park,

Chiang Mai Province, Thailand (Golovatch

& Enghoff 1993; Enghoff 2005).

3

Tylopus allorugosus Golovatch & Enghoff, 1993

Doi Inthanon National Park; Doi Suthep

National Park, Chiang Mai Province,

Thailand (Golovatch & Enghoff 1993;

Likhitrakarn et al. 2010).

4

Tylopus amicus Golovatch & Enghoff, 1993

Doi Pha Horn Pole National Park, Chiang

Mai Province, Thailand (Golovatch &

Enghoff 1993).

5

Tylopus asper Golovatch & Enghoff, 1993

Doi Inthanon National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993).

6

Tylopus baenzigeri Golovatch & Enghoff, 1993

Doi Suthep National Park (Golovatch &

Enghoff 1993); Royal Agricultural Station,

Angkhang Chiang Mai Province, Thailand

(see above).

7

Tylopus bispinosus Likhitralcarn, Golovatch,

Prateepasen & Panha, 2010

Umphang, Tale Province, Thailand

(Likhitrakarn et al. 2010).

8

Tylopus coriaceus Golovatch & Enghoff, 1993

Phu Kheio, Chaiyapum Province, Thailand

(Golovatch & Enghoff 1993).

9

Tylopus corrugatus Likhitrakarn, Golovatch &

Panha, 2014

Doi Inthanon National Park, Chiang Mai

Province, Thailand (Likhitrakarn et al.

2014a).

10

Tylopus crassipes Golovatch, 1984

Oquyho (Golovatch 1984); Nam Xay

commune; Hoang Lien National Park, Lao

cai Province, Vietnam (Nguyen 2012).

11

Tylopus degerboelae Golovatch & Enghoff, 1993

Doi Inthanon National Park; Doi Chiang

Dao National Park (Golovatch & Enghoff

1993); Doi Suthep National Park (Golovatch

& Enghoff 1993, Likhitrakarn et al. 2010);

Doi Phatang, Chiang Mai Province,

Thailand (Likhitrakarn et al. 2010).

12

Tylopus deharvengi Liu & Luo, 2013

Yaonan Dong, Guangxi, China (Liu & Luo

2013).

13

Tylopus doriae (Pocock, 1895)

Yado; Bia-po; Meteleo; Puepoli, Myanmar

(Pocock 1895); Doi Suthep National Park,

Chiang Mai Province, Thailand (Golovatch

& Enghoff 1993).

35

European Journal of Taxonomy 195: 1-47 (2016)

No.

Species

Localities

14

Tylopus dorsalis sp. nov.

Phuthalang forest protected area, Phongsali

Province, Laos (see above)

15

Tylopus extremus Likhitrakarn, Golovatch,

Prateepasen & Panha, 2010

Doi Phahom Pole National Park, Fang

District, Chiang Mai Province, Thailand

(Likhitrakarn et al. 2010).

16

Tylopusflavolineatas sp. nov.

Wat Tham Santisuk, Lopburi Province

Thailand (see above)

17

Tylopus golovatchi Nguyen, 2012

Xuan Son National Park, Phu Tho Province,

Vietnam (Nguyen 2012).

18

Tylopus grandis Likhitrakarn, Golovatch,

Prateepasen & Panha, 2010

Pha Mon Cave; Mae Lana, Pangmapha

District, Mae Hong Son Province

(Likhitrakarn et al. 2010).

19

Tylopus granulatus Golovatch, 1984

Cue Phuong Nature Reserve, Ninh binh

Province, Vietnam (Golovatch 1984).

20

Tylopus haplorugosus Golovatch & Enghoff,

1993

Doi Inthanon National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993, Likhitrakarn et al. 2010).

21

Tylopus hilaris (Attems, 1937)

Bana, Danang Province (Attems 1937);

Bach Ma National Park, Thua Thien Hue

Province; Ngoc Linh Mountain, Kon Turn

Province, Vietnam (Nguyen 2012).

22

Tylopus hilaroides Golovatch, 1984

Cue Phuong Nature Reserve Ninh binh

Province (Golovatch 1984); Cue Phuong

National Park, Ninh Binh Province,

Vietnam (Nguyen 2012).

23

Tylopus hoffmani Golovatch & Enghoff, 1993

Doi Suthep National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993).

24

Tylopus hongkhraiensis sp. nov.

Huai Hong Khrai Royal Development

Study Centre, Chiang Mai Province,

Thailand (see above)

25

Tylopus jeekeli Golovatch & Enghoff, 1993

Doi Inthanon National park (Golovatch &

Enghoff 1993); Doi Suthep National Park,

Chiang Mai Province, Thailand (Golovatch

& Enghoff 1993; Likhitrakarn et al. 2010).

26

Tylopus kabaki Golovatch, 2014

Deqen, Yunnan Province, China (Golovatch

2014).

27

Tylopus maculatus Golovatch, 1984

Oquyho, Lao cai Province, Vietnam

(Golovatch 1984).

28

Tylopus magicus Golovatch, 1984

Oquyho, Lao cai Province, Vietnam

(Golovatch 1984).

29

Tylopus moniliformis sp. nov.

Tad Fane Waterfall, Champasak Province,

Laos (see above)

30

Tylopus mutilatus (Attems, 1953)

Langbiang mountain, Lam Dong Province,

Vietnam (Attems 1953).

36

LIKHITRAKARN N. et al. , The millipede genus Tylopus

No.

Species

Localities

31

Tylopus nodulipes (Attems, 1953)

Luang Prabang, Laos; Mt Fan-Si-Pan

(Attems 1953); Nam Xay commune, Lao

Cai Province; Son Tay commune, Ha Tint)

Province; Pu Mat National Park, Nghe An

Province, Vietnam (Nguyen 2012).

32

Tylopus pallidus Golovatch & Enghoff, 1993

Doi Pha Horn Pole, Chiang Mai Province,

Thailand (Golovatch & Enghoff 1993).

33

Tylopus parahilaroides Likhitrakarn, Golovatch

& Panha, 2014

Phuluang Wildlife Sanctuary, Phuluang

District, Loei Province, Thailand

(Likhitrakarn et al. 2014a).

34

Tylopus parajeekeli Likhitrakarn, Golovatch,

Prateepasen & Panha, 2010

Doi Inthanon National Park, Chiang Mai

Province, Thailand (l ikhitrakarn et al. 2010).

35

Tylopus perarmatus Hoffman, 1973

Doi Suthep National Park (Hoffman 1973;

Golovatch & Enghoff 1993); Doi Inthanon

National Park (Golovatch & Enghoff 1993;

Likhitrakarn et al. 2010); Doi Chiang Dao

(Golovatch & Enghoff 1993); Doi Phatang,

Chiang Mai Province (Likhitrakarn et

al. 2010); Ban Huai Kaeo (Golovatch &

Enghoff 1993); Thum Pha Thai, Lampang

Province; Ban Pang Rim Kon; Phucheefah;

Doi Pha Tang, Chiang Rai Province; Nam

Min Waterfall, Phayao Province; Tham

Pha Nang Khoi, Phrae Province;Ton

Tong Waterfall, Nan Province, Thailand

(Likhitrakarn et al. 2010).

36

Tylopus perplexus Golovatch & Enghoff, 1993

Doi Pha Horn Pole, Chiang Mai Province,

Thailand (Golovatch & Enghoff 1993).

37

Tylopusphanluongi Nguyen, 2012

Ngoc Linh Mountain, Kon Tun Province,

Vietnam (Nguyen 2012).

38

Tylopus poolpermorum Golovatch & Enghoff,

1993

Doi Pha Horn Pok, Chiang Mai Province,

Thailand (Golovatch & Enghoff 1993).

39

Tylopus procurvus Golovatch, 1984

Oquyho (Golovatch 1984); Hoang Lien

National Park, Lao cai Province, Vietnam

(Nguyen 2012).

40

Tylopus prosperus Golovatch & Enghoff, 1993

Doi Inthanon National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993; Likhitrakarn et al. 2010).

41

Tylopus pulvinipes Golovatch & Enghoff, 1993

Phu Kheio, Chaiyaphum Province, Thailand

(Golovatch & Enghoff 1993).

42

Tylopus punctus sp. nov.

Mintaingbin Forest Camp, Chan State,

Myanmar (see above).

43

Tylopus re ductus Golovatch, 2013

Gaoligong Shan Mountain, Yunnan

Province, China (Golovatch 2013).

44

Tylopus retusus sp. nov.

Xay city, Oudomxai Province, Laos (see

below).

45

Tylopus roseiparaterga Nguyen, 2012

Tam Dao National Park, Vinh Phuc

Province, Vietnam (Nguyen 2012).

37

European Journal of Taxonomy 195: 1 —47 (2016)

No.

Species

Localities

46

Tylopus rugosus Golovatch & Enghoff, 1993

Chiang Dao (Golovatch & Enghoff, 1993);

Buathong Waterfall forest park, Chiang

Mai Province, Thailand (Likhitralcarn et al.

2010).

47

Tylopus sapaensis Nguyen, 2012

Hoang Lien National Park, Lao Cai

Province, Vietnam (Nguyen 2012).

48

Tylopus schawalleri Golovatch, 2013

Diancang Shan Mountain, Yunnan Province,

China (Golovatch 2013).

49

Tylopus semirugosus Golovatch & Enghoff, 1993

Ban Mussoe, Mae Sot District, Talc

Province, Thailand (Golovatch & Enghoff

1993).

50

Tylopus sigma (Attems, 1953)

Sapa, Lao Cai Province, Vietnam (Attems

1953).

51

Tylopus silvestris (Pocock, 1895)

village of Thao, Myanmar (Pocock 1895).

52

Tylopus similirugosus Golovatch & Enghoff,

1993

Doi Suthep National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993).

53

Tylopus similis Golovatch, 2014

Between Lijiang and Shangrila, Yunnan

Province, China (Golovatch 2014).

54

Tylopus sinensis Golovatch, 1995

Mengzi County, Yunnan Province, China

(Golovatch 1995).

55

Tylopus spinisterna Nguyen, 2012

Bi Doup-Nui Ba National Park, Lam Dong

Province, Vietnam (Nguyen 2012).

56

Tylopus strongylosomoides (Korsos &

Golovatch, 1989)

Tam Dao National Park, Vinh Phu Province

(Korsos & Golovatch 1989); Xuan Son

national Park, Phu Tho Province, Vietnam

(Nguyen 2012).

57

Tylopus subcoriaceus Golovatch & Enghoff,

1993

Doi Suthep National Park, Chiang Mai

Province, Thailand (Golovatch & Enghoff

1993).

58

Tylopus tamdaoensis Korsos & Golovatch, 1989

Tam Dao National Park, Vinh Phu Province,

Vietnam (Korsos & Golovatch 1989;

Nguyen 2012).

59

Tylopus thunghaihin sp. nov.

Plain of Jars, Xieng Khouang Province,

Laos (see above).

60

Tylopus topali Golovatch, 1984

Cue Phuong Nature Reserve, Ninh binh

Province (Golovatch 1984; Nguyen 2012);

Xuan Son National Park, Phu Tho Province,

Vietnam (Nguyen 2012).

61

Tylopus trigonum Likhitralcarn, Golovatch &

Panha, 2014

Pa Wai waterfall, Talc Province, Thailand

(Likhitralcarn et al. 2014a).

62

Tylopus veliger Likhitralcarn, Golovatch,

Prateepasen & Panha, 2010

Ton Tong Waterfall, Nan Province, Thailand

(Likhitralcarn et al. 2010).

38

LIKHITRAKARN N. et al. , The millipede genus Tylopus

Key to the known species of Tylopus (based chiefly on c?), modified after Likhitrakarn et al. (2010):

1. Most of S prefemora evidently swollen laterally (Figs 13C, 24C) .2

- All S prefemora normal, not bulged laterally (Figs 2G, 1 IF, G, 17E-G, 19C) .29

2. Surface of metaterga virtually smooth, at best extremely faintly rugulose in certain places and/

or with a few barely traceable (setigerous) tubercles near caudal margin (setae mostly broken off)

(Figs 7B, D, 12A-B, D, 21 A, C).3

- Surface of metaterga mostly rugulose to coarsely rugose/tuberculate (Figs 18A-B, D, 23B, D)

..9

3. Paraterga moderately developed (Figs 1A, C, 21 A, C), ratio of <$ midbody prozonite to metazonite

width below 1:1.2 .4

- Paraterga relatively well-developed (Figs 10B, D, 11B, D, 18B, D), radio of $ midbody

prozonite to metazonite width over 1:1.2 .5

4. Transverse sulcus on metaterga starting with segment 5. Calluses without incisions. Gonopod with

process hsmall and dentiform. Thailand.Golovatch&Enghoff, 1993

- Transverse sulcus on metaterga starting with segment 4. Calluses with faint one or two denticles

on poreless and poriferous paraterga, respectively. Gonopod very simple, without processes.

Southern China. T. reductus Golovatch, 2013

' China

Myanmar

-L.:-iP- '

Vietnam^-"

Thailand

Cambodia

Fig. 26. Distribution of the new eight species of Tylopus. Filled triangle. T. punctus sp. nov. Filled

circle. T. acuminatus sp. nov. Crossed circle. T. acuminatus sp. nov., T. dorsalis sp. nov. Crossed

square. T. acuminatus sp. nov. T. retusus sp. nov. Open circle. T. retusus sp. nov. Open triangle.

T. baenzigeri Golovatch & Enghoff, 1993. Filled square. T. hongkhraiensis sp. nov. Open diamond.

T. thunghaihin sp. nov. Inverted filled triangle. T. flavolineatus sp. nov. Open square. T. moniliformis

sp. nov.

39

European Journal of Taxonomy 195: 1-47 (2016)

5. Calluses without incisions. Gonopod postfemoral lobe I much broader than long; area basal to 1

delimited by a distinct cingulum.6

- Calluses mostly with 1-2 incisions. Gonopod postfemoral lobe 1 either as long as broad or longer;

no cingulum basal to 1.7

6. Body smaller: length up to ca 19-30 mm. Gonopod with a broadly coiled solenophore and a small

finger-shaped process on lobe 1. China . T. kabaki Golovatch, 2014

- Body larger: length ca 33-35 mm. Gonopod with a slender and coiled solenophore, but without

processes on lobe 1. Vietnam. T. magicus Golovatch, 1984

7. Metaterga at best only very faintly rugulose near stricture, near sulcus and/or at base of paraterga, with

2-3 weak, oblong tubercles near rear margin. S postfemora and tibiae tuberculiferous. Gonopods

with process z serrate, with 3 spines. Thailand. T. similirugosus Golovatch & Enghoff, 1993

- Metatergal surface entirely smooth, polished, without tubercles. $ legs without adenostyles

(= tubercles). Gonopods with process z pointed, without spines .8

8. Midline wanting. Gonopods with process h without additional tooth, while process z very high.

Vietnam. T. mutilatus (Attems, 1953)

- Midline mostly traceable on prozonae and anterior halves of metaterga. Gonopods with process h

higherwith an evident dorsal tooth, while process z small (Figs 13A-B, 14-15). Laos.

. T. acuminatus sp. nov.

9. Body larger: 37M2 mm long, 2.8-3.8 and 4.1-5.0 mm wide on pro- and metaterga, respectively.

.10

- Body smaller.15

10. Sterna between S coxae 4 with a pair of separated cones. Gonopod with a short spiniform

process h, a basally only poorly delimited lobe 1 and a small lobe-shaped process z. Thailand

. T. grandis Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

- Sterna between S coxae 4 with a slightly notched sternal lobe. Gonopod with a rather long process

h carrying two branches, plus a basally evidently delimited lobe 1 and a large, serrated process z.

Vietnam . T. phanluongi Nguyen, 2012

11. Both processes h and z of gonopod spiniform. Thailand.

. T. bispinosus Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

- Gonopod otherwise .12

12. Gonopod process h subflagelliform, process m extremely long and prominent. Thailand.

. T. extremus Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

- Gonopod otherwise .13

13. Gonopod process h prominent, spine s high, rather slender, curved, nearly pointed. Vietnam

. T. sapaensis Nguyen, 2012

- Gonopod otherwise .14

14. S legs shorter, ca 1.2-1.3 times as long as midbody height. Gonopod lobe 1 velum-shaped and

supplied with two denticles; spine z short and knife-shaped, while spine h rudimentary. Thailand

. T. veliger Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

- S legs longer, ca 1.6-1.7 times as long as midbody height. Gonopod spine z small, placed closer to base

of spine h. Thailand. T. parajeekeli Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

40

LIKHITRAKARN N. et al ., The millipede genus Tylopus

15. Most of metaterga with a pattern of 2+2 and 2+2 setigerous tubercles in two rows, rear row

somewhat less strongly developed than fore one. Myanmar and Thailand ... T. doriae (Pocock, 1895)

- Most of metaterga with rear row of setigerous tubercles or wrinkles more strongly developed than

fore row, the latter (next to) wanting.16

16. Transverse sulcus on metaterga starting with segment 4, either fully or almost fully developed there,

always fully developed starting with segment 5.17

- Transverse sulcus on metaterga starting only with segment 5.20

17. Transverse sulcus fully developed and reaching base of paraterga already starting with segment 4.

Gonopod tooth z at base of lobe 1 coarsely serrate along proximal margin. Vietnam.

. T. hilaris (Attems, 1937)

- Transverse sulcus fully developed only starting with segment 5. Gonopod tooth z either devoid of

serration or serrate along distal margin.18

18. Paraterga 2 rather broadly rounded caudolaterally. Gonopod relatively simple, process h poorly

developed, no additional outgrowths near base. Thailand .

. T. affinis Golovatch & Enghoff, 1993

- Paraterga 2 pointed caudally. Gonopods more complex.19

19. Coloration dark brown, without a cingulate pattern. Sternal lamina between S coxae 4 low

and distinctly bimodal. Gonopod tooth z prominent and serrate along distal margin. Thailand

. T. rugosus Golovatch & Enghoff, 1993

- Coloration pale, with a cingulate pattern. Sternal lamina between $ coxae 4 high, subquadrate.

Gonopod tooth z smaller and spiniform. Thailand. T. semirugosus Golovatch & Enghoff, 1993

20. Paratergal corner protruding caudad behind rear margin only starting with segment 15, being

obtuse-angled or subrectangular and lying more or less within the margin until segment 14.

Vietnam. T. hilaroides Golovatch, 1984

- Paratergal corner protruding caudad behind rear margin before segment 14, mostly pointed ...21

21. Pattern of tergal setation on segments 18 and/or 19: 2+2 and 5+5 in two rows .22

- Pattern of tergal setation at least on segments 5-19: 2+2 and 4+4 in two rows .25

22. Pattern of tergal setation 2+2 and 5+5 on both segments 18 and 19. Paraterga 2 pointed caudally.

Epiproct with pre-apical incisions very close to apical knobs. Sternal lamina between S coxae 4

an unusually low and even ridge. Adenostyles on midbody $ postfemora and, to a lesser extent,

tibiae exceptionally prominent. Thailand . T. poolpermorum Golovatch & Enghoff, 1993

- Pattern of tergal setation 2+2 and 5+5 on segment 19. Paraterga 2 more or less narrowly rounded.

Pre-apical incisions on epiproct more strongly removed from tip. Sternal lamina between S

coxae 4 concave medially. Ventral adenostyles on $ legs less prominent .23

23. Body smaller: width ca 2.0 mm. Sternal lamina between $ coxae 4 a pair of separated setigerous

tubercles. Ventral adenostyles on $ legs almost missing. Gonopods without any outgrowths near base

of process h. Thailand. T. haplorugosus Golovatch & Enghoff, 1993

- Body larger: width over 3.0 mm. Sternal lamina between $ coxae 4 single. Ventral adenostyles

on $ legs more prominent. Gonopod with a spine near base of process h.24

24. Sternal lamina between $ coxae 4 high, emarginated. Adenostyles on $ postfemora and tibiae

well-developed. Gonopods rather simple, spine z inconspicuous. Thailand .

. T. allorugosus Golovatch & Enghoff, 1993

41

European Journal of Taxonomy 195: 1 —47 (2016)

- Sternal lamina between <f coxae lower, slightly concave. Adenostyles on $ postfemora and

tibiae less strongly developed. Gonopods more complex, spine z long and large. Thailand

. T. perarmatus Hoffman, 1973

25. Paraterga 2 pointed caudally. Sternal lamina between S coxae 4 exceptionally densely setose, low

and concave ventrally. Gonopods with a medium-sized process h and a smaller lobular z at base of h.

Thailand. T. jeekeli Golovatch & Enghoff, 1993

- Paraterga 2 more or less narrowly rounded caudally. Sternal lamina between $ coxae 4 higher

and less strongly setose. Gonopod outgrowths h and z either almost wanting or very large ...26

26. Sternal lamina between $ coxae 4 with a straight ventral margin. Pleurosternal carinae poorly

developed, in $ slightly drawn behind rear margin only until segments 8-10 .27

- Sternal lamina between $ coxae 4 slightly concave ventrally. Pleurosternal carinae better

developed, in $ slightly drawn behind rear margin at least until segment 15 .28

27. Body smaller: width up to 3.1-3.2 mm. Mid-dorsal line very clear on both halves of metaterga.

Gonopods relatively simple, with both h and z almost wanting. Thailand.

. T. lioffmani Golovatch & Enghoff, 1993

- Body larger: width 4.0-5.3 mm. Mid-dorsal line not so well-developed at least on rear halves

of metaterga. Gonopods more complex, with both h and z very conspicuous (Figs 24A-B, 25).

Thailand . T. baenzigeri Golovatch & Enghoff, 1993

28. Metatergum 19 slightly rugulose posteriorly. Calluses on segment 2 with three, on following paraterga

with two, incisions. Gonopods extremely complex, with numerous spiniform outgrowths. Thailand

. T. perplexus Golovatch & Enghoff, 1993

- Metatergum 19 entirely smooth. Calluses with two or three incisions on poreless and poriferous

paraterga, respectively. Gonopod less strongly differentiated. Thailand.

. T. amicus Golovatch & Enghoff, 1993

29. Either most of $ sterna with an oblique tubercle or spine, or only anterior sterna with a small cone.

near coxae.30

- Neither spines nor tubercles on $ sterna.37

30. Calluses virtually devoid of incisions.31

- Calluses always at least with one lateral incision.32

31. $ coxae 4 with a pair of separated cones. Gonopod process h small, while solenophore slender and

short. China. T. sinensis Golovatch, 1995

- S coxae 4 with an entire, subtrapeziform, sternal lobe. Gonopod process h larger, while solenophore

stout and long. Vietnam. T. golovatchi Nguyen, 2012

32. A pair of small, fully separated, setose cones between S coxae 4 (Fig. 101—J). Gonopod with

process h rather short, slightly flattened, blunt (Fig. 11A-D). Laos . T. retusus sp. nov.

- An entire lobe between S coxae 4. Gonopod otherwise.33

33. Metaterga strongly rugose/tuberculate/granulate (Fig. 18A-F, H) .34

- Metaterga entirely smooth and polished, devoid of evident tubercles, at best extremely faintly

rugulose near transverse sulcus (Fig. 1A-F) .35

42

LIKHITRAKARN N. et al. , The millipede genus Tylopus

34. Calluses mostly with one lateral, clear incision. Pleurosternal carinae missing after segment 17.

Gonopod processes h, z, m and spine x present (Fig. 19A-B). Laos ...I thunghaihin sp. nov.

- Calluses with two lateral setigerous incisions. Pleurosternal carinae missing after segment 12.

Only gonopod process h present. Vietnam. T. spinisterna Nguyen, 2012

35. Only rear sternum on most of $ segments with a pair of small spines. Metaterga mostly with 2+2

and 3+3 setigerous tubercles in two transverse rows. Pleurosternal carinae missing after segment

9. Myanmar . T. silvestris (Pocock, 1895)

- Fore and rear sterna of most of S segments with a pair of small tubercles and spines, respectively.

Pleurosternal carinae present until segment 18.36

36. Metaterga with four transverse rows of very small setigerous tubercles or setae. S tarsal brushes

present until leg 6. $ femora 5 with a very strong, distoventral, densely pilose adenostyle (Fig. 2F).

Gonopod process h higher and pointed (Fig. 2A-D). (Fig. 26). Thailand. T. flavolineatus sp. nov.

- Metaterga with two transverse rows of small tubercles or spines, respectively S tarsal brushes missing.

S legs normal. Gonopod process h very small. Vietnam. T. maculatus Golovatch, 1984

37. Paraterga virtually missing (Figs 4A-H, 7A-H) .38

- Paraterga well- or strongly developed (Figs 1A, C, 10B, D, 12B, D, 16B, D, 18B, D, 23B, D) ...39

38. Antennae shorter (Fig. 4A), drawn behind only body segment 2 when stretched dorsally. A small,

rounded, densely setose, sternal cone between S coxae 4 (Fig. 4I-J). Goponod process h prominent,

while process z long and rather slender (Figs 5-6). Thailand . T. hongkhraiensis sp. nov.

- Antennae longer (Fig. 7A), reaching behind body segment 3. An entire, high, inverted funnel-shaped,

sternal lobe between each of $ coxae 3 and 4 (Fig. 7I-J). Goponod without processes (Figs 8-9). Laos

. T. moniliformis sp. nov.

39. Metaterga entirely smooth and polished, devoid of evident tubercles, at best extremely faintly

rugulose near transverse sulcus.40

- Metaterga rather clearly rugose/tuberculate/granulate, posterior row of setae at least partly borne on

tubercles.46

40. Transverse sulcus on metaterga starting with segment 3 or 4, but fully developed and reaching

bases of paraterga starting with segment 4 or 5. Gonopod process h large (Fig. 17A-D).41

- Transverse sulcus on metaterga starting only with segment 5. Gonopod process h small (Fig. 22)... 44

41. Gonopod with a distinct sharp process on lobe 1. Ventral adenostyles on S legs absent. Southern China

. T. similis Golovatch, 2014

- Gonopod without processes on lobe 1. Ventral adenostyles on S legs: a distal knob on femur, a

distomedial knob on postfemur, and a parabasal knob on both tibia and tarsus .42

42. Head wider than collum and subequal in width to segment 3. Gonopod process h at about

midlength with a strong ventral outgrowth, while solenophore without processes. Vietnam.

. T. procurvus Golovatch, 1984

- Head narrower than collum. Gonopod process h without outgrowths, while process x present on

solenophore .43

43. Antennae shorter (Fig. 16A), extending only behind body segment 2 when stretched dorsally.

S tarsal brushes present until segment 6. Gonopod process x longer, the latter’s tip rounded

dorsally (Fig. 17A-D). Laos . T. dorsalis sp. nov.

43

European Journal of Taxonomy 195: 1-47 (2016)

- Antennae longer, reaching until body segment 5. 3 tarsal brushes present on all segments. Gonopod

process x shorter, its tip pointed. Vietnam. T. crassipes Golovatch, 1984

44. Small species: width 1.6 mm. Only a single row of 2+2 tergal setae. Metaterga entirely smooth.

Adenostyles on 3 legs present. Vietnam. T. sigma (Attems, 1953)

- Body wider than 2.0 mm. Metaterga with two transverse rows of setae or traceable insertion

points. Metaterga very finely rugulose only near transverse sulcus. Adenostyles on 3 legs absent

or represented by pads in their stead .45

45. Paraterga broader: 3.1 mm wide. Pleurosternal carinae present until segment 13 or 14. Pads present

on 3 femora, postfemora, tibiae (all distally) and tarsi (almost entirely). Thailand.

. T. pulvinipes Golovatch & Enghoff, 1993

- Paraterga narrower: 2.1-2.5 mm wide. Pleurosternal carinae present until segment 17. 3 legs

without pads. Myanmar. T. punctus sp. nov.

46. Metatergal surface polished and smooth except for two rows of conspicuous tubercles .47

- Metaterga at least partly rugulose/rugose to granular; at most one row of tubercles.52

47. 3 legs without adenostyles (= tubercles). Gonopods with process h higher than solenophore,

strongly twisted, pointed apically. Southern China. T. schawalleri Golovatch, 2013

- 3 legs with adenostyles (= tubercles) at least on segment 6. Gonopods otherwise. Vietnam ...48

48. Paraterga very poorly developed, rounded, low, projecting slightly behind rear margin like small

knobs only on segments 18 and 19. Calluses virtually devoid of incisions. Transverse sulcus

on metaterga poorly developed, starting already with segment 2, although fully developed only

starting with segment 5. A paramedian pair of denticles between 3 coxae 5 behind a prominent,

subquadrate lamina between 3 coxae 4. Gonopod process h entirely missing. Thailand

. T. strongylosomoides (Korsos & Golovatch, 1989)

- Paraterga better developed, protruding behind rear margin at least starting with segment 5. Calluses

always at least with one lateral incision. Transverse sulcus starting only with segments 3-5.

Dentiform tubercles between 3 coxae 5 missing. Gonopod process h invariably present.49

49. Body larger: more than 2 5 mm long. 3 tarsal brushes notpresent on all legs. Gonopod lobe 1 normal... 50

- Body smaller: less than 18 mm long. 3 tarsal brushes present on all legs. Gonopod lobe 1 with a

spine apically.51

50. 3 tarsal brushes present on legs 1-15. Pleurosternal carinae present until segment 9. Gonopod

process h larger, finger-shaped. Vietnam. T. roseiparaterga Nguyen, 2012

- 3 tarsal brushes present on legs 5-30. Pleurosternal carinae present until segment 17. Gonopod

process h small, slender and pointed. Southern China. T. deharvengi Tiu & Tuo, 2013

51. Paraterga acutangular caudally and pointed beak-like already starting with collum. Tergal setigerous

tubercles: 3+3 and 5+5 on segments 16-19. Adenostyle pattern on 3 legs: a distal knob on femora

and a parabasal knob on most of postfemora, tibiae and tarsi. Gonopod process h large, lamellar and

sigmoid. Vietnam. T. granulatus Golovatch, 1984

- Paraterga acutangular caudally and pointed beak-like only starting with segment 4. 2+2 and 4+4

tergal setigerous tubercles on segments 16-19. Adenostyle pattern on 3 legs: a proximal finger-

shaped tubercle crowned with a bunch of setae only on femora 6, 8 and 9. Gonopod process h smaller,

spiniform. Vietnam. T. topali Golovatch, 1984

44

LIKHITRAKARN N. et al ., The millipede genus Tylopus

52. Transverse sulcus on metaterga starting and fully developed only with segment 5. Ventral

adenostyles present on all S podomeres except coxa. Vietnam and/or Taos .53

- Transverse sulcus on metaterga starting with segment 4, but fully developed only starting with

segment 5. Ventral tubercles only on some of $ telopoditomeres. Thailand .54

53. Metaterga rugulose also in front of transverse sulcus, at rear margin with several oblong tubercles.

Sternal lamina between $ coxae 4 a pair of setigerous knobs. Neither gonopod lobe m nor lobe 1 spini-

gerous. Vietnam. T. tamdaoensis Korsos & Golovatch, 1989

- Metaterga rugose only behind transverse sulcus, without evident tubercles at rear margin. Sternal

lamina between $ coxae 4 single. Both gonopod lobe m and lobe I crowned with a spine. Taos

and Vietnam. T. nodulipes (Attems, 1953)

54. Metaterga modestly rugulose only near transverse sulcus, posteriorly neither granular nor

microtuberculate. Calluses broad. Gonopod process h simple, high, never particularly coiled; lobe 1

very moderately serrate at apex.55

- Metaterga distinctly rugose-granular/microtuberculate even on fore halves. Calluses narrow.

Gonopod process h better developed and more strongly coiled; lobe 1 apically either bare or with

a digitiform outgrowth .56

55. Caudal corner of paraterga pointed starting with segment 3. Pleurosternal carinae particularly

well-developed, drawn behind rear margin until segment 16 or 17. Adenostyles often present on

S prefemora . T. degerboelae Golovatch & Enghoff, 1993

- Caudal corners of paraterga mainly narrowly rounded, pointed only starting with segment 15. Pleuro¬

sternal carinae less strongly developed. T. pallidus Golovatch & Enghoff, 1993

56. Coloration dark, brown. Sternal lamina between S coxae 4 a pair of separated setigerous tubercles

preceded by another pair of very small tubercles between coxae 3. Gonopod lobe 1 devoid of an

apical process. T. asper Golovatch & Enghoff, 1993

- Coloration uniformly pale. Sternal lamina between S coxae 4 single, not accompanied by additional

tubercles in front or behind. Gonopod lobe 1 with a strong, apical, finger-shaped process .57

57. Mostly 3+3 tubercles at rear margin of metaterga. Sternal lamina between S coxae 4 distinctly

emarginate. Larger adenostyles close to midlength on S postfemora and tibiae, femora with a

distal knob. Gonopod process h slenderer and shorter, apex of lobule m not developed into a spine

. T. subcoriaceus Golovatch & Enghoff, 1993

- Mostly 4+4 tubercles at rear margin of metaterga. Sternal lamina between S coxae trapeziform.

Larger adenostyles on both postfemora and tibiae more distal, femoral knob missing. Gonopod

process h unusually prominent, with a hook at base; apex of lobule m spiniform.

. T. coriaceus Golovatch & Enghoff, 1993

Discussion

At the moment, a total of 62 species of Tylopus are known from Indochina and the adjacent parts of

southern China and Myanmar, including eight new species described above. Most of the species (31,

or > 50%) come from Thailand, followed by Vietnam (18), Laos (6), southern China (6) and Myanmar

(3). Country endemism is close to 100%, as only T. doriae and T. nodulipes have been recorded from

two of the countries at once. There is little doubt, however, that the above outlined picture is still far

from final.

Having collected millipedes throughout Thailand and Laos, we can state that most of the Tylopus species

are quite rare and restricted in distribution, even though many occur sympatrically if not syntopically.

45

European Journal of Taxonomy 195: 1—47 (2016)

Thus, the Doi Inthanon and Doi Suthep mountains, both in northern Thailand, support at least 10

congeners each (Tikhitrakarn et al. 2014a). It seems that such sympatric assemblages are represented

by a number of phenofaunas separated by certain time periods rather than a geographic distance. As a

result, the adults of most of the known species have only been collected once and from a single locality,

with repeated collecting success being rather an exception than a rule.

An additional factor to be considered, the one that only makes the picture even more complicated, lies

in the fact that the bulk of Tylopus diversity appears to be confined to montane woodlands exceeding

500 m in elevation. This only exacerbates the situation as too many of such places are difficult to access

and thus only contribute to the abundant lacunae to be covered by future collecting efforts. Given the

highly mosaic, often relictual distribution of mountainous forests in the regions concerned, against the

background of too numerous undercollected to completely unsampled mountainous areas across most

of Indochina, as well as southern China and the East Indies, the real diversity of Tylopus can still be

very generously termed as highly underestimated. Thus, during the past two decades, since the first large

review of Tylopus which focused on the fauna of Thailand alone (see Golovatch & Enghoff 1993), the

number of described congeners known to occur in that country has nearly doubled, whereas the whole

generic diversity tripled. Yet Cambodia remains a complete terra incognita in this respect (Likhitrakarn

et al. 2015).

Acknowledgements

This project was partly funded through grants received from the Office of the Royal Development

Projects Board (RDPB), while most of the financial support was obtained from The Thailand Research

Fund, The TRF Senior Research Scholar RTA 5880002 (2015-2018) to SP. We thank the members of

the Animal Systematics Research Unit for their invaluable assistance in the field.

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Manuscript received: 30 November 2015

Manuscript accepted: 11 January 2016

Published on: 10 May 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

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, De nm ark

47