Corresponding authors: Sergei I. Golovatch (
Academic editor: R. Mesibov
The large genus
The genus
Ever since its proposal, the name
The present paper continues our studies on the still quite poorly known diplopod fauna of Southeast Asia (
Among the species groups currently delimited in
Unlike most of the genera of Polydesmida, including
So we are inclined to abandon species group delimitation in
Because an analysis of the phylogeny of either
Most of the older species revised, especially those valid ones in which the original descriptions appear to be deficient, are redescribed and illustrated here in sufficient detail to ensure their easy recognition.
New material derives from throughout Thailand and from northern Malaysia, taken from 1962 to 2011. Coloration was photographed in the laboratory (both live and alcohol material) for all of the encountered species. Material was then preserved in 75% ethanol and studied in the laboratory using an Olympus stereomicroscope. Scanning electron micrographs (SEM) were taken of uncoated specimens using a JEOL, JSM–5410 LV microscope. After SEM examination of the gonopods, they were returned to alcohol. Material of each of the species available for (re)study was photographed, the digital images assembled using the automontage software techniques, while gonopods also redrawn. Specimens were received from the following museum collections:
American Museum of Natural History, New York, USA,
Museum of Zoology, Chulalongkorn University, Bangkok, Thailand,
Natural History Museum, London, Great Britain,
Naturhistorisches Museum Wien, Austria,
Muséum d’histoire naturelle, Genève, Switzerland,
Zoological Museum, State University of Moscow, Russia,
Natural History Museum of Denmark, University of Copenhagen, Denmark.
So as not to repeat information, diagnoses are only provided here for new species, because the key below will show the main distinctions for all of the species in the genus.
In the catalogue sections, D stands for the original description, subsequent descriptive notes or appearance in a key, R for a subsequent record or records, N for giving a new name, and M for a mere mention. Not all of the relevant references are being quoted under certain Indochinese species, because there is no reason for duplicating the regional catalogues available and still valid for the millipedes of Vietnam and Thailand (
A dynamic web page for each taxon name mentioned in the paper is generated on the fly by the Pensoft Taxon Profile tool (see
A genus of
Gonopod with a long, subcylindrical, distodorsally usually setose coxite and a normal, cylindrical cannula. Telopodite mostly very slender and long, modestly curved. Prefemoral portion densely setose, about as long as (rarely) to ca 2–3 times (usually) shorter than femorite (measured together with “postfemoral” part lying distal to lateral sulcus). Femorite without evidence of torsion (= seminal groove running only mesally), often slightly enlarged distally, mostly with a clear-cut, oblique, distolateral sulcus demarcating a “postfemoral” part. Solenophore only moderately strongly curved mesad or caudomesad, consisting of modestly developed laminae lateralis and medialis, yet with lamina lateralis somewhat larger than lamina medialis, both sheathing a similarly long, simple, flagelliform solenomere with a barely exposed tip; tip of solenophore never deeply split, normally poorly bi- or trifid, some of apical prongs being either minute denticles or lappets, or small teeth, or completely reduced.
Type-species:
The
Based on the broad paraterga and the conformation of the solenophore,
Syntypes of
Length 33–38 mm (♂), 36–38 mm (♀), width of midbody pro- and metazona 2.7–2.8 and 4.0–4.2 mm (♂), 3.1–3.3 and 4.3–4.4 mm (♀), respectively. Coloration of alcohol material upon long-term preservation rather uniformly brown with contrasting pale yellowish paraterga, venter and legs light yellow-brown (
Head usual, clypeolabral region sparsely setose, surface of vertex smooth; epicranial suture distinct. Antennae moderately long (
Sterna sparsely setose, without modifications, but with a pair of small, rounded, completely separated, setose cones between ♂ coxae 4 (
Gonopods (
A complete historical review of the typification of
Originally described as a subspecies of
With the above synonymization, the nomenclature of
(
1 ♂, 2 ♀(CUMZ), Thailand, Chiang Mai Prov., Mae Rim Distr., Queen Sirikit Botanical Garden,
Length 14.5–20.5 (♂) or 16.5–27.5 mm (♀), width of midbody pro- and metazona 1.1–1.7 and 1.5–2.7 mm (♂), 1.1–2.5 and 1.6–3.2 mm (♀), respectively.Coloration, texture, all main somatic and gonopod characters as in
Gonopods (
This pantropical species has been described and redescribed several times, often under different names (e.g.
Our restudy of the type material of
♂ (NHMW-3500), Vietnam, Lamdong Prov., Dalat, 1500 m, 31.01.1935, leg. C. Dawydoff.
2 ♂, 1 ♀, 10 juv. (NHMW-3500), same locality, together with lectotype. Numerous fragments (NHMW-3499), Vietnam, Lamdong Prov., Peak Lang Biang, Arbre-Broyé, 1400 m, 02.02.1931, leg. C. Dawydoff.
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male.
Length 28–31 mm (♂), 34 mm (♀), width of midbody pro- and metazona 2.3–2.8 and 3.8–4.1 mm (♂), 3.4 and 4.6 mm (♀), respectively (vs up to 38 mm length, as given in the original descriptions (
Head usual, clypeolabral region sparsely setose (vs densely setose, as given in the original descriptions (
Sterna sparsely setose, without modifications, but with a very evident, high, setose, central cone between ♂ coxae 4 (
Gonopods (
This species is only known from southern Vietnam (Dalat and Peak Lang Biang).
♂ (CUMZ), Thailand, Phang Nga Prov., Khura Buri Distr., Similan National Park, Ko Bangu, Island 9, ca 40 m,
2 ♀ (CUMZ), same locality, together with holotype.
To emphasize the picturesque appearance of the animals.
The new species differs in the small size (up to 23 mm long and 3.5 mm wide), coupled with a particular coloration and a bifid gonopod tip (see also Key below).
Length ca 19 mm (♂) and 22–23 mm (♀), width of midbody pro- and metazona 1.8 and 2.7 mm (♂), 2.3–2.5 and 3.4–3.5 mm (♀), respectively. Coloration of live animals blackish, paraterga and epiproct contrasting creamy orange, legs and venter brownish to pale brown; coloration of alcohol material after preservation faded to castaneous brown or pale brown, paraterga (marbled at base) and epiproct somewhat faded to pale pinkish or pale yellow, legs and venter paler brown (
Clypeolabral region densely setose, vertex sparsely setose, epicranial suture distinct. Antennae moderately long, clavate (antennomere 7 broad) (
Head in width < collum < segments 3–4 < 2 = 5–16 (♂, ♀); thereafter body gently and gradually tapering. Collum with three transverse rows of setae, 4+4 anterior, 2+2 intermediate, and 2+2 posterior; caudal corner of paraterga very narrowly rounded, slightly bordered and declined ventrally, not extending behind tergal margin (
Sterna delicately and sparsely setose, without modifications; a paramedian pair of small, strongly separated, setose tubercles between ♂ coxae 4. Gonopod aperture broken during removal of gonopods. Legs moderately long and slender, slightly incrassate in ♂, midbody ones ca 1.2–1.4 (♂) or 0.9–1.1 times (♀) as long as body height, prefemora without modifications, ♂ tarsal brushes present until legs of segment 8.
Gonopods (
Among all nine islets of the Similan Archipelago, Thailand inspected for millipedes in April 2010, only three appeared to support
♂ (CUMZ), Thailand, Nakhon Ratchasima Prov., Pakchong Distr., Khao Rup Chang, ca 420 m,
2 ♂, 2 ♀ (ZMUC), 2 ♂, 2 ♀ (ZMUM), 14 ♂, 21 ♀ (CUMZ), same data as holotype. 1 ♂ (CUMZ), same Distr., Klang Dong Restaurant, ca 360 m,
To emphasize the evident metatergal tuberculation.
The new species differs in evidently tuberculated metaterga, coupled with only a small anterolateral incision on paraterga and a bifid gonopod tip (see also Key below).
Length 24–32 mm (♂), 27–34 mm (♀), width of midbody pro- and metazona 2.6–3.3 and 4.5–5.2 mm (♂), 2.9–4.0 and 4.5–5.5 mm (♀), respectively Coloration of live animals blackish, paraterga and epiproct contrasting creamy orange, legs and venter brownish to pale brown (
Clypeolabral region densely setose, vertex bare, epicranial suture distinct. Antennae short, poorly clavate (
Head in width << collum < segments 3–4 < 2 = 5–16 (♂), or head < collum < segments 3–4 < 2 < 5–16 (♀); thereafter body gently and gradually tapering. Collum with three transverse rows of medium-sized setae, 4+4 anterior, 2+2 intermediate, and 2+2 posterior, all borne on very evident tubercle, these being especially high in caudal row; paraterga slightly declivous, broadly rounded and narrowly bordered, caudal corner a minute knob, not extending behind tergal margin (
Sterna delicately and rather densely setose; a paramedian pair of evident, fully separated, setose cones between ♂ coxae 4 (
Gonopods (
There are two further populations which we regard as representing the same new species. One of the populations (15 ♂, 3 ♀ (CUZM), non-types) comes from Thailand, Lop Buri Prov., Khok Samrong Distr., Khao Wong Phrachan Temple,
Length 22–30 mm (♂), 24–31 mm (♀), width of midbody pro- and metazona 2.6–2.9 and 3.9–4.2 mm (♂), 2.8–3.3 and 4.0–4.3 mm (♀), respectively. Colour pattern same as in the type series of
All other characters as in the typical
Venter and legs sometimes as pallid as paraterga and epiproct. Paraterga slightly less prominent, narrower (
Length 36 mm (♂), 36–37 mm (♀), width of midbody pro- and metazona 3.0 and 4.9 mm (♂), 3.5–3.7 and 5.2–5.5 mm (♀), respectively. Colour pattern same as in the type series of
All other characters as in the typical
Paraterga less prominent, even midbody ones nearly not produced behind tergal margin (
♂ (CUMZ), Thailand, Nakhon Ratchasima Prov., Wang Nam Khiao Distr., Sakaerat Enviromental Research Station,
1 ♂ (CUMZ), same data as holotype. 1 ♂, 3 ♀ (CUMZ), same locality, 10.01.2007, leg. N. Likhitrakarn.
To emphasize the strong similarity to
Comes closest to
Length 21–23 mm (♂), 23–24 mm (♀), width of midbody pro- and metazona 2.2–2.4 and 3.5–4.2 mm (♂), 2.1–3.7 and 3.4–4.3 mm (♀), respectively. Coloration of alcohol material after preservation uniformly blackish-brown (♀) or apparently faded to uniformly brown (♂), with paraterga, venter, distal part of epiproct and several basal podomeres more flavous, pallid to light yellow (
All other characters as in
Antennae longer (
Head in width < collum < segments 3–4 < 2 < 5–16 (♂, ♀); thereafter body gently and gradually tapering. Paraterga on collum slightly declivous, subtriangular, with a small, but evident indentation near midway and a small, caudally directed, sharp denticle at caudal corner, the latter not extending behind tergal margin (
Sterna delicately and sparsely setose. Only small paramedian knobs in front of gonopod aperture. Legs moderately long and slender, almost not incrassate in ♂, midbody ones ca 1.2–1.3 (♂) or 0.9–1.1 times (♀) as long as body height, prefemora without modifications.
Gonopods (
This new species has been found rather close to the localities of
♂ (CUMZ), Thailand, Surin Prov., Mueang Surin Distr., Khao Phanom Sawai National Park, ca 200 m,
1 ♂, 1 ♀ (ZMUC), 1 ♂, 1 ♀ (ZMUM), 3 ♀ (CUMZ), same data, together with holotype. 2 ♂ (CUMZ), Prachinburi Prov., Prachantakham Distr., Takror Waterfall, ca 30 m,
To emphasize this species being quite common in the eastern part of Thailand close to the border with Cambodia.
Differs in unequal terminal lobes of the solenophore, both of which show a minute tooth near their bases, coupled with pointed, subtriangular paraterga on the collum etc. (see also Key below).
Length 31–38 (♂) or 32–38 mm (♀), width of midbody pro- and metazona 2.6–4.0 and 4.2–4.7 mm (♂), 3.2–3.8 and 4.8–5.3 mm (♀), respectively.
Coloration of live animals (
Clypeolabral region densely setose, vertex sparsely setose, epicranial suture distinct. Antennae moderately long, clavate (antennomere 6 broadest), extending behind body segment 2 (♂) (
Sterna sparsely setose, without modifications; cross-impressions shallow; with a paramedian pair of very small, flat, strongly separated, setose knobs between ♂ coxae 4 (
Gonopods (
This new species appears to be quite widespread in the eastern part of Thailand close to the border with Cambodia (
♂ (CUMZ), Thailand, Chanthaburi Prov., Khlung Distr., Troknong Waterfall,
25 ♂, 30 ♀ (CUMZ), 3 ♂, 3 ♀ (ZMUC), 3 ♂, 3 ♀ (ZMUM), same data, together with holotype.
To emphasize this species being not quite typical to readily fit into any of the former species groups.
Superficially, this new species strongly resembles
Length 32–39.5 (♂) or 34.5–44 mm (♀), width of midbody pro- and metazona 3.0–3.6 and 5.0–5.6 mm (♂), 3.6–4.0 and 5.5–6.4 mm (♀), respectively.
Coloration of live animals (
Clypeolabral region densely setose, vertex barely setose, epicranial suture distinct. Antennae moderately long (
Sterna sparsely setose, without modifications; cross-impressions shallow, especially so due to a superficial axial impression; a large, setose, transverse lobe bearing a paramedian pair of large, basally contiguous cones between ♂ coxae 4 (
Gonopods (
♂ (CUMZ), Thailand, Chanthaburi Prov., Pong Nam Ron Distr., Hin Dard Waterfall, ca 260 m,
To emphasize the extremely broad paraterga.
Differs in the extremely broad paraterga, coupled with the pleurosternal carinae represented by complete high crests with a sharp caudal tooth on segments 2–7 (♂) etc. (see also Key below).
Length 31 mm, width of midbody pro- and metazona 3.1 and 5.0 mm, respectively.
Live coloration (
Clypeolabral region sparsely setose, vertex bare, epicranial suture distinct. Antennae moderately long (
Sterna sparsely setose, without modifications; cross-impressions shallow; lobe between ♂ coxae 4 much like in
Gonopods (
This new species shows the paraterga relatively perhaps among the broadest amongst congeners.
♂ (CUMZ), Thailand, Nong Bua Lamphu Prov., Suwannakhuha Distr., near Suwannakhuha Cave,
1 ♀ (CUMZ), same data, together with holotype.
To emphasize the suberect gonopod femorite.
Differs in a suberect gonofemorite, coupled with mostly flavous metaterga and widely separated sternal cones between ♂ coxae 4 (see also Key below).
Length ca 26 (♂) or 28 mm (♀), width of midbody pro- and metazona 2.5 and 3.9 mm (♂), 3.0 and 4.3 mm (♀), respectively.
Coloration of alcohol material after long-term preservation castaneous brown with a pattern of contrasting whitish to light brown paraterga and epiproct, and mostly greyish-white to light brownish posterior halves of postcollum metaterga, with caudal edges and front 1/4 of metaterga, as well as surface below paraterga and entire rear halves of prozona brown to dark brown; head and antennomeres 6 and 7 brown to dark brown; venter and a few basal podomeres light brownish to brown, legs growing infuscate (brown) distally (
Clypeolabral region sparsely setose, vertex bare, epicranial suture distinct. Antennae moderately long, clavate (antennomere 6 broadest), extending behind body segment 2 (♂, ♀) dorsally. Head in width < collum < segments 2 < 3 < 4 < 5–17 (♂), or head < segment 3–4 < 2 < 5–17 (♀); thereafter body gently and gradually tapering. Collum with three transverse rows of setae: 4+4 anterior, 2+2 intermediate and 4+4 posterior; paraterga slightly declivous, broadly rounded and narrowly bordered, caudal corner a small knob not extending behind tergal margin (
Sterna sparsely setose, without modifications; cross-impressions shallow; a paramedian pair of evident, small, fully separated, setose cones between ♂ coxae 4 (
Gonopods (
This species has long been acknowledged as being quite disjunct due to the gonopod showing a parabasally twisted femorite, while the tip is rather deeply bifid, somewhat intermediate between that of typical
♂ (NHMW-3507), Indonesia, East Java Prov., Lumajang Regency, shore of Ranu (= lake) Bedali, 10.10.1928, leg. A. F. Thienemann & H. J. Feuerborn.
3 ♂, 3 ♀ (NHMW-8004), Vietnam, Khanh Hoa Prov., Cua Bé (4–6 km from Nha Trang), no date, det. C. Attems. 2 ♂, 1 ♀ (NHMW-8005), same Prov., Nha Trang, no date, det. Attems. 4 ♂, 4 ♀ (NHMW-8007), Vietnam, Tinh Soc Trang (? or Bà Ria-Vũng Tàu), Poulo Condore Island, no date, leg. C. Dawydoff, det. C. Attems. 4 ♂, 1 ♀ (NHMW-8006), Cambodia, Kampot Prov., Bokor Mt in Damrei Mountains (= Elephant Mountains), near Bok Kor Village, no date, leg. C. Dawydoff, det. C. Attems. 4 ♂, 1 ♀ (NHMW-8008), Indonesia, West Java Prov., Tjisaroea, leg. W. S. S. van Benthem-Jutting, det. C. Attems. 7 ♂, 8 ♀, 2 juv. (NHMW-8009), Indonesia, East Java Prov., Bondowoso, Ijen Caldera, no date, det. C. Attems.
Length 17–30 mm (♂), 19–29 mm (♀), width of midbody pro- and metazona 1.4–2.2 and 2.0–3.1 mm (♂), 1.6–2.7 and 2.0–3.4 mm (♀), respectively (vs 3.3 mm in holotype, up to 4.0 mm in width, as given in the available descriptions (
Head usual, clypeolabral region densely setose, surface of vertex smooth (vs a pair of strong setae, as given in the descriptions (
Sterna sparsely setose, without modifications, but with two small, low, rounded, fully separated, setose knobs between ♂ coxae 4 (
Gonopods (
This species was originally described in two varieties, each based on a single male:
♂ (NHMW-3508), Indonesia, central Java, Sarangan, 09.12.1928, leg. A. F. Thienemann & H. J. Feuerborn.
Length ca 29 mm, width of midbody pro- and metazona 2.2 and 3.3 mm, respectively. Coloration of alcohol material upon long-term preservation rather uniformly brown (
All other characters as in
Collum with three transverse rows of setae, 4+4 anterior, 2+2 intermediate, and 2+2 posterior setae; caudal corner of paraterga subrectangular, narrowly rounded (
Sterna sparsely setose, with neither modifications nor knobs between ♂ coxae 4 (
Gonopods (
This variety was elevated to a full species by (
♂ of
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male coming from a certain locality, because (1)
Length 41 mm, width of midbody pro- and metazona 3.1 and 4.4 mm, respectively (vs 38 mm in length and 4.6 mm in width, as given in the original description (
Head usual, clypeolabral region sparsely setose, surface of vertex smooth, epicranial suture very distinct. Antennae rather long and slender (
Sterna densely setose, without modifications, but with two large, rounded, fully separated, setose cones between ♂ coxae 4 (
Gonopods (
This species was renamed to avoid homonymy (
♂ of
1 ♂ of
1 ♀ of
2 ♂, 1 ♀, 1 juv. of
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male coming from a certain locality, because (1) (
Length 29–31 mm (♂), ca 31–35 mm (♀), width of midbody pro- and metazona 2.5–3.0 and 3.7–4.1 mm (♂), 3.4 and 4.5 mm (♀), respectively (vs 2.5 and 4.0 mm in width, as given in the available descriptions (
Head usual, clypeolabral region densely setose, surface of vertex smooth, epicranial suture distinct. Antennae rather long and slender (
Sterna sparsely setose, without modifications, but with two rather large, rounded, fully separated, but nearly contiguous, setose cones between ♂ coxae 4 (
Gonopods (
This species is known from Lamdong Province, southern Vietnam and Luang Prabang Province, Laos (
The above is a combined redescription, based on material of both nominate species.
♂ (NHMW-3501), Cambodia, Kampot Prov., Ream, Sre-Umbell, 1938–39, leg. C. Dawydoff.
1 ♀ (NHMW-3501), same locality, together with lectotype.
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male.
Length ca 31 mm (♂), 32 mm (♀), width of midbody pro- and metazona 2.2 and 3.4 mm (♂), 3.1 and 4.3 mm (♀), respectively (vs 3.2–4.0 mm in width, as given in the original description (
Head usual, clypeolabral region densely setose, surface of vertex smooth, with 1+1 setae flanking a distinct epicranial suture. Antennae rather long and slender (
Sterna sparsely setose, without modifications, but with two large, rounded, fully separated, setose cones between ♂ coxae 4 (
Gonopods (
♂ (NHMW-3518), Indonesia, eastern Java, Tengger Mountains, no date, leg. H. Fruhsdorfer.
1 ♂, 6 ♀ (NHMW-7998); 1 ♂, 1 ♀ (NHMW-7999); 8 ♂, 15 ♀ (NHMW-8003), Indonesia, Java, Tjibodas, 1897, leg. H. A. Möller, det. Attems. 4 ♂, 1 ♀ (NHMW-8002), Indonesia, Java, no date, leg. T. H. Adensamer, det. C. Attems.
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male, because the type series was stated to have been shared between the collections of the Berlin and Vienna museums (
Length 24–28 mm (♂), 29–33 mm (♀), width of midbody pro- and metazona 2.6–2.8 and 3.4–4.1 mm (♂), 3.3–3.7 and 4.5–5.1 mm (♀), respectively (vs 40 mm in length and 4.0 mm in width, as given in the available descriptions (
Head usual, clypeolabral region sparsely setose, surface of vertex smooth and bare, epicranial suture distinct. Antennae rather long and slender (
Sterna sparsely setose, without modifications, but with two small, rounded, fully separated, but subcontiguous, setose cones between ♂ coxae 4 (
Gonopods (
♂ (NHMW-3506), Vietnam, Hon Ba Island, Nhatrang, 06.1930, leg. C. Dawydoff.
1 ♀ (NHMW-3506), same locality, together with lectotype. 1 ♀ (NHMW-3505), Vietnam, Darlac, frontier du Cambodge, 07.1930, leg. C. Dawydoff.
Lectotype designation proposed herewith is necessary to ensure the species is based on a complete male.
Length ca 38 mm (lectotype), 31–34 mm (♀), width of midbody pro- and metazona 3.4 and 5.0 mm (lectotype), 2.9–3.4 and 4.2–4.4 mm (♀), respectively (vs 3.0 and 5.0 in width, as given in the available descriptions (
Head usual, clypeolabral region densely setose, surface of vertex smooth, with a few setae flanking a distinct epicranial suture. Antennae long and slender (
Sterna sparsely setose, without modifications, but with a large, central, setose cone between ♂ coxae 4 (
Gonopods (
2 ♂, 1 ♀ (MHNG), Indonesia, Java, no date, leg. L. Zehntner.
Length 19–22 (♂) or 24 mm (♀), width of midbody pro- and metazona 1.4–1.5 and 2.1–2.4 mm (♂), 2.0 and 2.6 mm (♀), respectively (vs 2.8 mm in width, as given in the original description (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae moderately long, clavate (antennomere 6 broadest), extending behind body segment 3 (♂) or 2 (♀) dorsally (
Sterna sparsely setose, without modifications; cross-impressions shallow; with a paramedian pair of very evident, anteroventrally directed lobe deeply, but incompletely split into two between ♂ coxae 4 (
Gonopods (
The above redescription is meant to augment the fairly complete one by
♂ (MHNG), Malaysia, environs of Kuala Lumpur, Batu Caves, 24.02.1975, leg. P. Strinati.
Length 39 mm, width of midbody pro- and metazona 2.9 and 4.4 mm, respectively. Coloration of alcohol material upon long-term preservation dark castaneous brown with contrasting yellowish paraterga; venter, legs yellownish and tip of epiproct light red-brown, antennae also light red-brown, but antenomere 6 slightly infuscate, brownish, antenomere 7 dark brown, and tip of antennae pallid (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae rather short, clavate (antennomere 6 broadest), extending behind body segment 2 dorsally. Head in width < collum < segment 2 < 3 < 4 < 5–16; thereafter body gently and gradually tapering. Collum with three transverse rows of setae, 4+4 anterior, 2+2 intermediate, and 3+3 posterior; caudal corner of paraterga nearly pointed, slightly declined ventrally and continuing collum’s convexity (
Sterna sparsely setose, without modifications; cross-impressions shallow; a pair of strongly separated, anteroventrally directed, narrowly rounded, setose cones between ♂ coxae 4 (
Gonopods (
The original description is fairly complete, also illustrated in due detail (
♂(NHML), Singapore, Mile 8, Old Upper Thomson Road, 30.12.1962, leg. Kok Oi Yee.
1 ♂, 5 ♀ (CUMZ), Singapore, Central Region, Nature Reserve, Bukit Timah,
Length 34–35 (♂) or 3.3–3.6 mm (♀), width of midbody pro- and metazona 2.5–2.6 and 3.9–4.4 mm (♂), 3.4–3.6, and 4.8–4.9 mm (♀), respectively.
Coloration of live animals (
The original description of this species was so complete and detailed (
♂(NHMW-3509), Indonesia, Sumatra, Lampung Prov., Way Kambas, forest road, ca 10 m, 26.12.1993, leg. R. Melisch.
The original description of this species was so complete and detailed (
♂(ZMUC), Thailand, Krabi Prov., road between Krabi and Phuket, 10 km north of Krabi,
1 ♂ (CUMZ), Thailand, Phuket Prov., Mueang Distr., Kathu Waterfall, ca 10 m,
Length 31–35 (♂), width of midbody pro- and metazona 2.2–3.0 and 4.0–4.6 mm (♂), respectively (vs 34–37 mm in length, 2.2–3.4 and 4.0–4.7 mm in width on midbody pro- and metazona, respectively, both in ♂ and ♀, as given in the original description (
Coloration of live animals (
The original description of this species was so complete and detailed (
This species occurs, so far as known, only on Phuket Island and in the adjacent Krabi Province, southern Thailand.
♂(ZMUC), Thailand, Trang Prov., Hat Chao Mai National Park,
1 ♂ (CUMZ), Thailand, Nakhon Si Thammarat Prov., Nopphitam Distr., Krungshing Waterfall,
Length 27–36 (♂) or 30–38 mm (♀), width of midbody pro- and metazona 2.1–3.5 and 3.6–4.5 mm (♂), 2.7–3.5 and 4.0–5.1 mm (♀), respectively.
Coloration of live animals (
The original description of this species was so complete and detailed (
This species appears to be quite widespread in the southern parts of Thailand (
♂ (ZMUC), Thailand, Phuket Prov., Mueang Phuket Distr., Ton Sai Waterfall,
2 ♂, 1 ♀ (CUMZ), Thailand, Chumphon Prov., Lang Suan Distr., Khao Kriab Temple, ca 100 m,
Length 33–42 (♂) or 33–51 mm (♀), width of midbody pro- and metazona 2.6–3.6 and 4.0–6.0 mm (♂), 2.9–4.5 and 4.7–6.7 mm (♀), respectively.
Coloration of live animals (
Calluses of several midbody and/or posterior metaterga (16–19) can be less strongly convex laterally (
Tip of solenophore narrowly rounded to sharply dentiform, rather depending on aspect (
The original description of this species was complete and detailed enough (
♂(ZMUC), Thailand, Yala Prov., Ban Lang National Park,
Coloration of alcohol material upon long-term preservation probably faded to dark castanceous brown with contrasting pallid paraterga, middle part of prozona, epiproct, venter and pleurosternal carinae (
The original description of this species was so complete and detailed (
♂(ZMUC), Thailand, Satun Prov., Thale Ban National Park,
1 ♀ (CUMZ), Thailand, Satun Prov., Khuan Don Distr., Thale Ban National Park, ca 110 m,
Length 36–42 (♂) or 39–46 mm (♀), width of midbody pro- and metazona 2.8–3.6 and 4.9–5.6 mm (♂), 3.6–4.1 and 5.3–6.0 mm (♀), respectively.
Coloration of live animals (
The original description of this species was so complete and detailed (
♂(ZMUC), Thailand, Krabi Prov., road between Krabi and Phuket, 10 km north of Krabi,
2 ♂, 2 ♀ (ZMUC), 2 ♂, 2 ♀ (ZMUM), 15 ♂, 16 ♀ (CUMZ), Thailand, Surat Thani Prov., Kanchanadit Distr., Tham Khao Phanomwang,
Length 35–49 (♂) or 35–52 mm (♀), width of midbody pro- and metazona 3.1–4.1 and 5.0–6.5 mm (♂), 3.4–4.8 and 5.0–7.1 mm (♀), respectively.
Coloration of live animals (
Collum with caudal corner of paraterga ranging from obtusangular (described as being about 100˚ in holotype,
Metatergal sulcus visible on segments 5–18, more rarely incomplete also on segment 19 (holotype). Pleurosternal carinae expressed as complete high crests with a sharp caudal tooth on segment 2, thereafter increasingly well divided into a front bulge and a caudal tooth, the latter sharp or rounded, on segments 2–7; both front bulge and caudal tooth increasingly strongly reduced either until segment 10 (11), with a small sharp tooth until segment 16 (17) (♂), or expressed as complete high crests with a sharp caudal tooth on segments 2–4, thereafter increasingly well divided into a front bulge and a caudal tooth, sharp or rounded, on segments 5–10, a small sharp tooth until segment 16 (17) (♀). Midbody legs ca 1.1–1.3 (♂) or 0.9–1.2 times (♀) as long as body height.
Base of lamina lateralis with a small, but conspicuous ventral knob (like in
This species appears to be widespread in southern Thailand (
2 ♀ (CUMZ), Thailand, Mae Hong Son Prov., Mueang Mae Hong Son Distr., Tham Woa Temple,
Length 33–41.5 mm (♂) to 34–51 mm (♀), width of midbody pro- and metazona 3.1–4.0 and 4.9–5.7 mm (♂), 3.2–4.5 and 5.2–7.0 mm (♀), respectively.
Coloration of live animals (
Antennae rather long, clavate (antennomere 6 broadest), extending behind body segment 3 (♂) or 2 (♀) dorsally.
Collum with caudal corner of paraterga ranging from obtusangular (
Midbody legs ca 1.1–1.4 (♂) to 0.9–1.3 times (♀) as long as body height.
Tip of gonopod trifid, with terminal lobule longest, middle spiniform prong sometimes being shorter than usual (
This species appears to be widespread over much of Thailand, as well as in the adjacent parts of Myanmar (
This species still remains dubious (
This species has been beautifully redescribed by
This species seems to be endemic to southern Myanmar together with the adjacent Mergui Archipelago and Thailand.
This species has been adequately redescribed and illustrated from a ♂ syntype by
This species has been adequately redescribed and illustrated from a ♂ syntype by
This species has been adequately redescribed and illustrated from the ♀ lectotype by
This species seems to have been described from a single ♂ (holotype) (
This species has been redescribed and illustrated from the ♂ lectotype coming from Gunong Inas, Perak State and two ♀ paralectotypes stemming from Kuala Aring, Kelantan State, Malaysia (
This species has been described from the ♂ holotype (
This species has recently been redescribed in due detail from the type series taken from Wat Son (Son Temple), near Bandon River, Kanchanadit District, Surat Thani Province, Thailand (
This species has been adequately described from Kebun Raya, Bali, Indonesia (
This species has been adequately described from Pass Puncak, 34 km from Bogor, Java, Indonesia (
This species has recently been redescribed both from the type series and new material coming from the northwestern coast of the Gulf of Siam, Thailand (
This species has recently been described from several places in west-central Thailand (
This species has recently been described from several places in southern Thailand and the adjacent parts of Malaysia (
This species has recently been described from several places in northern and central Thailand (
This species has recently been described from a single locality in the central part of Malay Peninsula within Thailand (
♂ (CUMZ), Thailand, Srakaeo Prov., Khao Chakan Distr., Khao Sam Sip,
2 ♂, 2 ♀ (ZMUC), 2 ♂, 2 ♀ (ZMUM), 4 ♂, 13 ♀, 2 juv. (CUMZ), same locality, together with holotype. 3 ♂, 1 ♀ (CUMZ), Chonburi Prov., Mueang Chon Buri Distr., Ang Hin Station,
Honours to Dr. Kumpol Isarankura, Professor of the Department of Biology of Chulalongkorn University, who collected some of the type specimens.
Differs in a peculiar colour pattern, coupled with narrow calluses of paraterga, and very broadly separated cones between ♂ coxae 4 (see also Key below).
Length mostly 29–34 (♂) or 33–36.5 mm (♀), width of midbody pro- and metazona 2.45–2.75 and 3.8–4.35 mm (♂), 2.9–3.35 and 4.35–4.85 mm (♀), respectively. One ♂ paratype (from Krathing Waterfall), 39.5 mm long, 3.6 and 5.5 mm wide on midbody pro- and metazona, respectively.
Coloration of alcohol material after long-term preservation castaneous brown with a pattern of contrasting whitish to yellow paraterga and epiproct, and mostly greyish-white posterior halves of postcollum metaterga; head and antennomeres 6 and 7 brown to dark brown; venter and a few basal podomeres light brown to yellow-brown, legs growing infuscate (brown) distally; tip of antenna pallid (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae moderately long (
Sterna delicately and sparsely setose, without modifications, but with a pair of very small, blunt, fully separated cones between ♂ coxae 4 (
Gonopods (
This new species is rather widespread in eastern Thailand (
♂ (CUMZ), Malaysia, Perak State, Bondig, Air Banum, ca 500 m,
2 ♀ (CUMZ), same data, together with holotype. 1 ♀ (CUMZ), Malaysia, Johor State, Sungai Bantang, ca 100 m,
To emphasize the mostly strongly elevated paraterga.
Differs by mostly strongly elevated paraterga, coupled with two small sternal cones between ♂ coxae 4 (see also Key below).
Length 34 (holotype) or 31–38 mm (♀), width of midbody pro- and metazona 2.8 and 4.3 mm (holotype), 2.9–3.5 and 4.3–5.0 mm (♀), respectively.
Coloration of live material (
Clypeolabral region densely setose, vertex bare, epicranial suture distinct. Antennae short, slightly clavate (antennomere 6 broadest), extending behind body segment 3 (♂) or 2 (♀) dorsally. Head in width < collum < segments 3 and 4 < 2 < 5–17 (♂, ♀); thereafter body gently and gradually tapering. Collum with three transverse rows of medium-sized setae, pattern untraceable; paraterga slightly declivous, subtriangular, with a faint lateral incision near midway, evidently discontinuing dorsum’s convexity, caudal corner pointed, about level to rear tergal margin (
Sterna sparsely setose, shining, without modifications; cross-impressions shallow; a paramedian pair of rather small, but evident tubercles in front of gonopod aperture; a small central lobe with a paramedian pair of evident, setose, apical cones between ♂ coxae 4 (
Gonopods (
This new species is rather widespread in northern Malaysia (
♂ (CUMZ), Malaysia, Kedah State, Gunung Baling,
2 ♀ (CUMZ), same data, together with holotype. 1 ♂ (CUMZ), Malaysia, Perlis State, Kaki Bukit,
To emphasize the strong similarity to
Comes closest to
Length 40–42 (♂) or 41–45 mm (♀), width of midbody pro- and metazona 3.0–3.7 and 4.0–5.6 mm (♂), 3.0–4.2 and 4.9–6.1 mm (♀), respectively. Holotype 3.95 mm long, 3.9 and 4.0 mm wide on midbody pro- and metazona, respectively.
Coloration of alcohol material after long-term preservation brown to dark brown with contrasting whitish calluses, lateral parts of paraterga close to calluses and, sometimes, entire epiproct, as well as light yellow to yellow-brown legs and epiproct; two paramedian spots divided by a narrow, brown or grey-brown axial stripe on prozona and central parts of metaterga slightly lighter than background, light brown, except for an infuscate cross composed of transverse sulcus and axial line, the latter often extending from collum to tip of epiproct (
All other characters as in
Antennae barely reaching end of segment 3 (♂) or 2 (♀) dorsally. Ribs at caudal margin of metaterga often vague, sometimes represented by low bosses. Axial line always visible on collum, as well as on following prozona, metaterga and epiproct. Paraterga very strongly developed (
Sterna densely setose. Legs long and slender, slightly incrassate in ♂, midbody ones ca 1.7–1.8 (♂) or 1.2–1.3 times (♀) as long as body height; ♂ tarsal brushes visible on legs 1–8, thereafter absent.
Gonopods (
This new species is rather widespread in northern Malaysia (
♂ (CUMZ), Thailand, Phang Nga Prov., Khura Buri Distr., Similan National Park, Ko Miang, Island 4, ca 40 m,
1 ♂, 1 ♀ (ZMUC), 1 ♂, 1 ♀ (ZMUM), 4 ♂, 3 ♀ (CUMZ), same locality, together with holotype.
To emphasize the type locality.
Differs by a strongly contrasting colour pattern, coupled with pleurosternal carinae represented by complete crests bulged anteriorly and with a sharp caudal tooth on segments 2–7, thereafter only a sharp caudal tooth on segments 8–16 (♂), or crests bulged anteriorly and with a sharp caudal tooth on segments 2–4, thereafter only a small sharp caudal tooth on segments 5–14 (♀) (see also Key below).
Length 26–30 (♂) or 31–33 mm (♀), width of midbody pro- and metazona 2.1–2.6 and 3.5–3.8 mm (♂), 2.9–3.4 and 4.0–4.3 mm (♀), respectively.
Coloration of live animals blackish, paraterga and epiproct contrastingly creamy yellow, legs and venter dark brown to blackish; coloration of alcohol material after long-term preservation dark brown to blackish, paraterga (marbled at base) and epiproct faded to pale yellow, legs and venter more pale brown (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae moderately long (
Sterna delicately and sparsely setose, without modifications, but with a pair of very small, blunt, fully separated cones between ♂ coxae 4 (
Gonopods (
This new species is only known from one of the islets of Similan Archipelago, Thailand (
♂ (CUMZ), Malaysia, Pahang State, Panang National Park, ca 80 m,
1 ♂, 1 ♀ (CUMZ), same State, Lake Kenyir, ca 170 m, 4°50'44"N, 102° 43'15"E, 25.05.2011, leg. S. Panha.
To emphasize the spiniform paraterga.
Differs in strongly spiniform paraterga, coupled with virtually smooth metaterga etc. (see also Key below).
Length 25–28 mm (♂), 29 mm (♀), width of midbody pro- and metazona 2.0–2.2 and 3.1–3.5 mm (♂), 2.4 and 3.5 mm (♀), respectively.
Coloration of live animals blackish-brown, calluses of paraterga (especially caudal parts) and distal half of epiproct contrasting red-brown; legs and venter yellow to light brown (
Clypeolabral region densely setose, vertex bare, epicranial suture distinct. Antennae short, poorly clavate (
Sterna shining, sparsely setose; a paramedian pair of evident, fully separated, anteroventrally directed, setose cones between ♂ coxae 4 (
Gonopods (
This new species seems to be restricted to Pahang State, southern Malay Peninsula within Malaysia (
♂ of
Length 25.5 mm, width of midbody pro- and metazona 2.2 and 3.2 mm, respectively.
Coloration of alcohol material after long-term preservation yellowish-brown with contrasting pallid paraterga; venter, legs and antennae light yellowish (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae moderately long, clavate (antennomere 6 broadest), extending behind body segment 3 dorsally (
Sterna sparsely setose, without modifications; cross-impressions shallow; with an evident, paramedian, sparsely setose, directed anteroventrally, bulge between ♂ coxae 4, this bulge showing traces of axial division (
Gonopods (
The original description of
Concerning the status of
3 ♀ (NHMW-7985), Indonesia, eastern Java, “Tanger region” (= Tengger Mountains), 1893–1894, leg. D. Adensamer.
Length 26–30 mm, width of midbody pro- and metazona 3.2–3.5 and 3.8–4.4 mm, respectively (vs 30 mm in length and 3.2 mm in width, as given in the original description (
Clypeolabral region densely setose, vertex sparsely so, epicranial suture distinct. Antennae rather long (antennomere 6 broadest), extending behind body segment 3 dorsally. Head in width < collum < segment 3 = 4 < 2 < 5–16; thereafter body gently and gradually tapering. Collum with three transverse rows of setae traceable only as insertion points: 3+3 anterior, 2+2 intermediate and 3+3 posterior; caudal corner of paraterga rounded, strongly declined ventrally and almost continuing collum’s convexity (
Sterna sparsely setose, without modifications; cross-impressions rather deep. Legs moderately long and slender, midbody ones ca 1.1–1.2 times as long as body height.
This species has been described from a series of ♀ syntypes, still known only from the type locality: Tengger Mountains, eastern Java, Indonesia (
In addition to
2 ♀ (NHMW-7986), Seychelles, Mahé Island, primeval forest, 1895, leg. A. Brauer.
Length 19–20 mm, width of midbody pro- and metazona 2.1–2.2 and 2.5–2.6 mm, respectively. Coloration of alcohol material upon long-term preservation uniformly light grey-brown (
Metaterga densely and irregularly setose. All other somatic characters as in
This species has been described from three ♀ syntypes, still known only from the type locality: Mahé Island, Seychelles (
Based both on the morphological characters (e.g. the narrow paraterga and densely setose metaterga) and distribution, there can be no doubt that
The paraterga in
The paraterga in
The paraterga in
According to the original description (
that of a species of the mainly Papuan tribe
A genus of
Gonopod with a long subcylindrical coxite and a usual, cylindrical cannula. Telopodite long and slender, modestly curved. Prefemoral part densely setose, more than 2 times shorter than femorite. The latter without evidence of torsion, slightly enlarged distally, without sulcus demarcating a “postfemoral” part. Solenophore consisting of subequally modestly developed laminae lateralis, and medialis, both sheathing a similarly long, simple, flagelliform solenomere with a barely exposed tip; tip of solenophore never deeply split, poorly bi- or trifid, at least some of its apical prongs being short spines.
Other species included:
1 ♂, 1 ♀ (NHMW-3517), Vietnam, Lamdong Prov., Dalat, 1500 m, 02.1933, leg. C. Dawydoff.
1 ♂ (NHMW-7987), Vietnam, Lamdong Prov., Dalat, Peak Lang Biang, 1200–2400 m, no date, leg. C. Dawydoff, det. C. Attems. This sample was erroneously labeled as a syntype, which cannot be such, because only material from Dalat was referred to in the available descriptions (
Length ca 18 mm (♂) or 20 mm (♀), width of midbody pro- and metazona 2.0 and 2.5 mm (♂), 2.2 and 2.8 mm (♀), respectively.
Coloration of alcohol material after long-term preservation faded to uniformly light brown (
Clypeolabral region densely setose, vertex bare, epicranial suture distinct. Antennae short, poorly clavate (
Sterna sparsely setose; a large central lobe with a paramedian pair of evident, subcontiguous, anteroventrally directed, setose cones between ♂ coxae 4 (
Gonopods (
1 | Gonopod tip like a single, very small, rounded lobule ( |
|
– | Gonopod tip mainly bi- to trifid. Modifications between ♂ coxae 4 mostly present. Local in Southeast Asia | 2 |
2 | Gonopod femorite about as long as prefemoral portion ( |
|
– | Gonopod femorite ca 2–3 times longer than prefemoral portion (e.g. |
3 |
3 | No distinct lateral sulcus on gonopod femorite demarcating a “postfemoral” region, at most a very faint one; solenophore tip neither acuminate nor branching. Thailand and northernmost Malaysia | 4 |
– | A distinct lateral sulcus on gonopod femorite demarcating a “postfemoral” region; solenophore tip either acuminate or rather evidently branching (e.g. |
9 |
4 | At least surface of metaterga leathery, dull | 5 |
– | Dorsal surface shining | 7 |
5 | Only caudal parts of paraterga contrasting light. Paraterga less strongly developed, narrower, with only a small incision at best in front 1/3 extent. No tergal setae borne on evident knobs/tubercles |
|
– | Entire paraterga contrastingly light, much broader, largely with an evident, dentiform incision in front 1/3 extent. At least some tergal setae borne on evident knobs/tubercles | 6 |
6 | Surface of metaterga rough, microgranulate and rugulose. Metatergal setigerous tubercles higher, present also on collum. Paratergal incisions especially prominent |
|
– | Surface of metaterga nearly smooth, microgranulate. Metatergal setigerous tubercles evidently smaller, absent from collum. Paratergal incisions evident, but not so prominent |
|
7 | Adults > 30 mm long and > 4 mm wide. Collum broadly rounded laterally. Paraterga set high, at least some upturned above dorsum. Sternal cones between ♂coxae 4 isolated. Lateral sulcus on gonofemorite traceable, but very faint |
|
– | Adults < 30 mm long and < 4 mm wide. Collum narrowly rounded to angulate laterally. Paraterga set high, but remaining below dorsum even in male. Sternal cones between ♂ coxae 4 fused basally into a single lamina. Lateral sulcus on gonofemorite absent | 8 |
8 | Bases of paraterga more broadly light. Legs shorter, midbody ones ca 1.2 (♂) or 1.0 times (♀) as long as body height. Tip of solenophore like an elongate and undulate lobe |
|
– | Bases of paraterga more narrowly light. Legs longer, midbody ones ca 1.5 (♂) or 1.2–1.3 times (♀) as long as body height. Tip of solenophore tridentate |
|
9 | Gonopod tip bifid ( |
10 |
– | Gonopod tip trifid, albeit middle prong can be very small ( |
18 |
10 | Metaterga smooth and shining, at most faintly rugulose ( |
11 |
– | Metaterga at least with one rather evident, transverse row of tubercles near caudal margin ( |
13 |
11 | Gonopod femorite slightly, but evidently twisted near base; tip rather deeply split, both terminal and subterminal prongs spiniform |
|
– | Gonopod femorite not twisted near base; tip shallowly bifid, both terminal and subterminal prongs dentiform (e.g. |
12 |
12 | Length ca 19–23 mm, width of midbody pro- and metazona 1.8 and 2.7 mm (♂), 2.3–2.5 and 3.4–3.5 mm (♀), respectively. Coloration of paraterga and epiproct creamy orange (live material) to faded pinkish or pale yellow (alcohol-preserved material) ( |
|
– | Length 31–38 mm, width of midbody pro- and metazona 2.6–4.0 and 4.2–4.7 mm (♂), 3.2–3.8 and 4.8–5.3 mm (♀), respectively. Coloration of paraterga and epiproct creamy yellow (both live and alcohol-preserved material) ( |
|
13 | Metaterga with only a single row of very small tubercle near caudal margin ( |
14 |
– | Metaterga with two rows of tubercles: small ones in front of, larger ones behind sulcus (near caudal margin) ( |
16 |
14 | Caudal corners of anterior and midbody paraterga not extended behind tergal margin ( |
|
– | Caudal corners of all postcollum paraterga evidently produced behind tergal margin. Metaterga mostly infuscate. Sternal cones between ♂ coxae 4 subcontiguous | 15 |
15 | Pleurosternal carinae complete high crests with a sharp caudal tooth on segments 2–7 (♂). Transverse sulcus visible on metaterga 5–18, narrow, not reaching bases of paraterga, ribbed at bottom. Stricture between pro- and metazona narrow and rather shallow, faintly beaded at bottom down to base of paraterga ( |
|
– | Pleurosternal carinae complete high crests with a sharp caudal tooth on segments 2–4(5) (♂, ♀). Transverse sulcus evident ( |
|
16 | A single sternal cone between ♂ coxae 4 ( |
|
– | Two sternal cones between ♂ coxae 4 ( |
17 |
17 | Postcollum paraterga with a very strong front indentation laterally ( |
|
– | Postcollum paraterga with only a very small front indentation laterally ( |
|
18 | Paraterga mostly level to or even above dorsum | 19 |
– | All paraterga below dorsum | 28 |
19 | Colour pattern not strikingly contrasting, calluses only being inconspicuously paler than a dark remaining background ( |
20 |
– | Coloration of metaterga more or less strongly contrasting, with very pale calluses against a very dark remaining background ( |
21 |
20 | Metaterga with two transverse rows of evident tubercles ( |
|
– | Metaterga quite smooth, without transverse rows of evident tubercles ( |
|
21 | Calluses on metaterga rather thin ( |
22 |
– | Calluses on metaterga rather thick ( |
23 |
22 | Sternal cone between ♂ coxae 4 single, large ( |
|
– | Two small sternal cones between ♂ coxae 4 ( |
|
23 | Metaterga virtually smooth, at most faintly rugulose, only seldom ( |
24 |
– | Metaterga with a caudal row of evident tubercles | 26 |
24 | ♂ tarsal brushes entirely wanting. Epiproct with very distinct apical papillae. Lombok Island, Indonesia |
|
– | ♂ tarsal brushes present. Epiproct with small apical papillae. Java Island, Indonesia | 25 |
25 | Tarsal brushes present until ♂ midbody legs |
|
– | Tarsal brushes present until ♂ legs 7 |
|
26 | Adult body particularly small, only ca 2.6 mm in width (♂). Bali Island, Indonesia |
|
– | Adult body larger, at least 3.5 mm in width. Java Island, Indonesia | 27 |
27 | Paraterga relatively strongly produced caudally, paratergal calluses in dorsal view rather strongly sinuate near ozopores |
|
– | Paraterga relatively slightly produced caudally, paratergal calluses in dorsal view rather poorly sinuate near ozopores |
|
28 | A row of very small tubercles near caudal margin of some metaterga ( |
29 |
– | Metaterga virtually smooth, at best only slightly rugulose near caudal margin ( |
31 |
29 | Small tubercles present only on metaterga 10–18(19) |
|
– | Small tubercles present on all postcollum metaterga ( |
30 |
30 | Paraterga more strongly produced caudally ( |
|
– | Paraterga less strongly produced caudally ( |
|
31 | A single sternal lobe between ♂ coxae 41 ( |
32 |
– | Two sternal cones, sometimes very low ones, between ♂ coxae 4 ( |
33 |
32 | Sternal lobe between ♂ coxae 4 subquadrate. Perak State, southern Malaysia |
|
– | Sternal lobe between ♂ coxae 4 divided into two lobules only apically ( |
|
33 | Metaterga and collum usually more or less flavous, with lighter bands, spots or markings, at least in caudal parts ( |
34 |
– | Metaterga and collum between paraterga usually completely dark ( |
41 |
34 | Pore-bearing midbody paraterga particularly thick in lateral view ( |
35 |
– | Pore-bearing midbody paraterga considerably thinner in lateral view ( |
36 |
35 | Paraterga set mostly slightly higher ( |
|
– | Paraterga set mostly slightly lower ( |
|
36 | Entire collum yellow, colour pattern like in |
|
– | Collum and subsequent tergites between paraterga at least partly infuscate, only postsulcus halves of metaterga often pallid to pale brown | 37 |
37 | No dark axial stripe on metaterga. Paraterga less prominent, their anterolateral margin relatively narrowly rounded ( |
38 |
– | A dark axial stripe or line on metaterga evident. Paraterga much more prominent, their anterolateral margin very broadly rounded ( |
39 |
38 | Colour pattern as in |
|
– | Colour pattern as in |
|
39 | Sternal cones between ♂ coxae 4 smaller ( |
|
– | Sternal cones between ♂ coxae 4 larger ( |
40 |
40 | Colour pattern of live animals as in |
|
– | Colour pattern of live animals as in |
|
41 | Postcollum metaterga flavous behind transverse sulcus ( |
42 |
– | Postcollum metaterga dark on both pre-sulcus and postsulcus halves ( |
45 |
42 | Paraterga with anterolateral corner considerably more angulate, more narrowly rounded |
|
– | Paraterga with anterolateral corner considerably more broadly rounded ( |
43 |
43 | Metaterga nearly completely flavous, only a thin axial line and transverse sulcus forming an infuscate cross. King Island, Mergui Archipelago, Myanmar |
|
– | Metaterga mostly dark, including bases of paraterga ( |
44 |
44 | Pleurosternal carinae present at least as a caudal denticle until segment 12 (♂). Tarsal brushes present until about ♂ midbody legs |
|
– | Pleurosternal carinae present at least as a caudal denticle until segment 16 (♂) or 13 (♀). Tarsal brushes present until about ♂ legs 8 | |
45 | Caudal corner of midbody calluses only modestly produced behind tergal margin ( |
46 |
– | Caudal corner of midbody calluses very strongly produced behind tergal margin ( |
47 |
46 | Pleurosternal carinae complete crests on segments 2–4 (♂, ♀) ( |
|
– | Pleurosternal carinae complete crests bulged anteriorly and with a sharp caudal tooth on segments 2–7, thereafter only a sharp caudal tooth on segments 8–16 (♂), or crests bulged anteriorly and with a sharp caudal tooth on segments 2–4, thereafter only a small sharp caudal tooth on segments 5–14 (♀). Tarsal brushes present until legs of ♂ segment 13 | |
47 | Caudal corner of paraterga evidently rounded ( |
|
– | Caudal corner of paraterga evidently pointed ( |
48 |
48 | Paraterga especially strongly produced behind tergal margin ( |
|
– | Paraterga much less strongly produced behind tergal margin ( |
49 |
49 | Transverse metatergal sulcus present only on segments 5–18 ( |
|
– | An incomplete transverse metatergal sulcus traceable also either on segments 3 and 4 or only on segment 4 | 50 |
50 | Transverse metatergal sulcus traceable on segments 3 and 4, fully developed on segments 5–18 ( |
|
– | Transverse metatergal sulcus incomplete only on segment 4, complete on segments 5–18 ( |
|
Distribution of the genus
Distributions of the new species of
Distributions of three widespread species of
Distribution of two widespread species of
The genus
There can be no doubt that further collecting efforts, especially in still very poorly explored regions as Laos, Myanmar and Cambodia, also some parts of Vietnam, will reveal many more species of
This project was funded through grants received from the Higher Education Research Promotion and National Research University Project of Thailand, Office of the Higher Education FW646A (2011–2013), and from the Chulalongkorn University under the Strategic Scholarship for Frontier Research Network for the Joint Ph.D. Program Thai Doctoral degree to SP. The authors especially SP would like to express the sincere Thanks to the National Science and Technology Development Agency (NSTDA) for financial support for flatback millipede studies in Thailand from the beginning to this publication. We are greatly obliged to Janet Beccaloni (NHML), Henrik Enghoff (ZMUC), Lorenzo Prendini (AMNH), Peter Schwendinger (MHNG) and Verena Stagl (NHMW) for kindly loaning us type material under their care. Thanks are also extended to members of the Animal Systematics Research Unit, Department of Biology, Faculty of Science for their help in collecting specimens.
Special thanks go to Robert Mesibov, Australia and an anonymous reviewer who have greatly improved the paper by spotting numerous errors and providing several valuable suggestions.