Revision of the Crematogaster ranavalonae-group in Asia , with description of two new species ( Hymenoptera , Formicidae )

The Asian members of the Crematogaster ranavalonae-group are revised and twelve species, including two new species, C. hashimi sp. n. and C. imperfecta sp. n. are recognized. The members are distinguished from the other Asian Crematogaster in having smooth, shiny bodies with short appressed setae on the surface. Crematogaster sikkimensis Forel, 1904 is raised to the species level, and the following new synonyms are established: C. aberrans Forel, 1892 = C. aberrans assmuthi Forel, 1913, syn. n. = C. aberrans inglebyi Forel, 1902, syn. n. = C. soror Forel, 1902, syn. n.; C. ebenina Forel, 1902 = C. ebenina corax Forel, 1902, syn. n. A key to the species based on the worker caste is provided.


Introduction
The hyperdiverse ant genus Crematogaster had been subdivided into sixteen subgenera (Emery 1922;Wheeler 1922). Recently a molecular phylogenetic analysis has suggested that most of these subgenera are non-monophyletic, and that the subgenus Oxygyne is monophyletic (Blaimer 2012c). Subsequently Blaimer (2012b)

Methods
Most observations were made on an Olympus SZX12 stereomicroscope. The scanning electron micrographs (SEM) were prepared with a JSM-5600LV scanning electron microscope. Color images were taken using a Canon EOS 50D with a Canon MP-E 65 mm 1-5× macro lens, then processed using Combine ZM. The relative position of propodeal spiracles was observed in lateral view, with the mesosoma carefully tilted to the position with the true maximum of Weber's length.

Measurements and indices
Measurements were made under an Olympus SZX12 stereomicroscope using micrometers. All measurements are expressed in millimeters, recorded to the second decimal place. The measurements for petiole and postpetiole follow Longino (2003).
Head Width (HW): Maximum width of head in full-face view, excluding the eyes (Fig. 1).
Head Length (HL): Perpendicular distance from vertex margin to line tangent to anteriormost projections of clypeus in full-face view (Fig. 1).
Weber's Length of the mesosoma (WL): Diagonal length, measured in lateral view from the anterior margin of the pronotum (excluding the collar) to the posterior extremity of the propodeal lobe (Fig. 4).
Petiole Length (  An important character in the Asian Crematogaster ranavalonae-group is explained below. Ventrolateral katepisternal ridge (Fig. 7). The ridge separates the mesopleuron lateral surface from ventral surface. Most Asian Crematogaster species have a well-defined ridge separating the two surfaces (Fig. 7B), but in some species the ridge is absent or vestigial (Fig. 7A). The ridge is visible in pinned specimens from lateral or ventrolateral view.

Asian Crematogaster ranavalonae-group
The Asian members of the Crematogaster ranavalonae-group not only have the morphological features diagnosed by Blaimer (2012aBlaimer ( , 2012b, but also show the additional features below. (i) Pronotum steeply raised in lateral view. (ii) Ventrolateral katepisternal ridge indistinct, but weakly developed anteriorly in some species. (iii) Integument essentially smooth and shiny. (iv) Erect pilosity almost absent. Some erect setae are developed on the clypeus or dorsal surface of petiole and postpetiole, but absent on the dorsum of head, mesosoma and fourth to seventh abdominal tergites. (v) Dorsum of head, mesosoma and fourth abdominal tergite with short and appressed setae.
This species group is easily defined from other Asian Crematogaster in having a steeply raised pronotum, smooth and shiny body surface, and short and appressed body setae.
Mesonotum highly convex in lateral view. Ventrolateral katepisternal ridge indistinct. Propodeal spine short and stout; the length variable, but generally as long as propodeal spiracle, dorsum not higher than anterior propodeum in lateral view. Pro- podeal spiracle as large as mesothoracic spiracle pit, situated apart from the propodeal declivity in lateral view, directed laterally.
In dorsal view, shape of petiole scoop with convex side, as long as broad. Petiole with anterolateral corner angulate. Petiolar spiracle big, as wide as half of propodeal spiracle in diameter, directed laterally. Postpetiole bilobed, with slight longitudinal median sulcus. Postpetiole slightly wider than petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus generally smooth and shiny, but weakly striated with faint rugulae. Malar region with feeble longitudinal rugulae. Dorsal surface of promesonotum weakly punctuated. Lateral surface of pronotum shiny, but weakly punctuated. Mesopleuron weakly punctuated. Dorsal surface of propodeum smooth and shiny. Lateral propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed, sparse setae. Clypeus without pair of longer setae on anteriormost portion. Anterior clypeal margin with one single longer seta and one pair of longer setae on median portion, mixed with some shorter setae on side. One to three pairs of short, erect setae on pronotal shoulder. Posterolateral tubercle with two pairs of erect setae. Ventral surface of petiole with longer appressed setae. Postpetiole with two pairs of longer setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown.

Distribution. This species is distributed in India and Thailand.
Comments. This species is unique among Asian Crematogaster ranavalonae-group in having the scape with sparse setae and smaller propodeal spiracles, situated apart from the propodeal declivity.
The syntype workers of C. aberrans assmuthi match well with syntype workers of C. aberrans. I treated C. aberrans assmuthi as a junior synonym of C. aberrans. The syntype workers of C. aberrans inglebyi match well with syntype workers of C. aberrans, but with only the following small difference: the former has right brown colored body (cf. Forel 1902). I concluded that C. aberrans inglebyi is a junior synonym of C. aberrans. The syntype workers of C. soror match well with syntype workers of C. aberrans. In Forel's original description (Forel 1902), he mentioned that the head is subquadratic. However, the posterior margin of head is not angulate, appearing rounded in front view. Thus I here synonymize C. soror with C. aberrans.

Crematogaster augusti Emery
Crematogaster augusti Emery, 1895: 31, pl. 2, figs. 19, 20.  Comments. Emery's original description was based on a single queen. Our collections are limited to worker specimens, but two unique features are found in the type. First, the propodeal spines are not developed, as they are in the queen of C. agnetis Forel from Madagascar (Blaimer 2012a). Second, the body surface appearing smooth and shiny. The taxonomic status will remain uncertain until nest series become available. General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is as long as width of scape. Compound eye large and slightly projecting beyond lateral margin of head in full face view.

Crematogaster butteli Forel
Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine long and stout, longer than spiracle diameter, directed upward, dorsum higher than anterior propodeum in lateral view. Propodeal spiracle large, situated on lateral surface of propodeum and close to propodeal declivity in lateral view.
In dorsal view, shape of petiole scoop with tapering side posteriorly, longer than broad. Petiole with anterolateral corner angulate. Petiolar spiracle big, as wide as half of propodeal spiracle in size. Postpetiole bilobed but without longitudinal median sulcus. Postpetiole as wide as petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region generally smooth, with feeble rugulae. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron shiny, but with longitudinal rugulae. Dorsal surface of propodeum smooth and shiny. Lateral propodeum shiny, but with longitudinal rugulae.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with sparse, short and appressed setae. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two pairs of longer setae on median portion, mixed with some shorter setae on sides. No erect setae on pronotal shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole with appressed setae. Postpetiole with one pair of longer erect setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown. Distribution. This species is known only from the type locality in Sumatra.
Comments. This species is similar to C. tumidula but can be distiguished in having angulate anterolateral corners of the petiole .  Fig. 28 Crematogaster (Oxygyne) daisyi Forel, 1901: 376   General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is as long as width of scape. Compound eye large and slightly projecting beyond lateral margin of head in full face view.
In dorsal view, shape of petiole scoop with convex side, longer than broad. Petiole with subparallel side on anterior half in dorsal view. Petiolar spiracle large, as large as half of propodeal spiracle, directed downward. Postpetiole bilobed with longitudinal median sulcus. Postpetiole wider than petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar regions generally smooth. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum generally shiny, but with feable rugulae. Dorsal surface of propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed setae sparsely. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two pairs of longer setae, mixed with some shorter setae on sides. No erect setae on pronotal shoulder. Posterolateral tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown. Distribution. This species is only known from Malaysia (Borneo). Comments. This species is similar to C. sikkimensis, but differs in having the spiracles on the petiole located on the lower position in lateral view. Fig. 29 Crematogaster (Oxygyne) dalyi Forel, 1902: 201  teeth smaller. Scape exceeding posterior corner of head, with dense appressed setae, each of which is less than width of scape. Compound eye slightly projecting beyond lateral margin of head in full face view.

Crematogaster dalyi Forel
Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine short and stout; length longer than spiracle, directed upward; dorsum as high as anterior propodeum in lateral view. Propodeal spiracle situated close to propodeal declivity in lateral view, directed laterally.
In dorsal view, shape of petiole scoop, longer than broad. Petiolar spiracle big, as wide as half of propodeal spiracle in size. Postpetiolar spiracle big and distinct, located anteriorly on lateral surface. In dorsal view, postpetiole broader than long, strongly bilobed but without longitudinal median sulcus. Postpetiole distinctly wider than petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus generally smooth and shiny, but with feable rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum generally smooth and shiny, but with longitudinal rugulae on anterior portion. Mesopleuron and lateral propodeum generally smooth and shiny, but with weak longitudinal rugulae. Dorsal surface of propodeum smooth and shiny.
Erect pilosity absent. Dorsum of head, clypeus and mesosoma with short and appressed setae. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with one single longer setae and two pairs of longer setae on the median portion, mixed with some shorter setae on sides. No erect seate on pronotal shoulders. Posterolateral tubercle without appressed setae. Ventral surface of petiole with appressed setae. Postpetiole without setae posteriorly. Fourth abdominal tergite with short and appressed setae.
Body color brown. Distribution. This species is only known from the type locality in India. Comments. This species is similar to C. sikkimensis, but differs in having short propodeal spines. Fig. 30 Crematogaster (Oxygyne) ebenina Forel, 1902: 199  teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is less than or as long as width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view.

Crematogaster ebenina Forel
Ventrolateral katepisternal ridge indistinct. Propodeal spine long and stout; length equal to slightly greater than diameter of propodeal spiracle, directed upward, dorsum as high as anterior propodeum. Propodeal spiracle large and situated close to propodeal declivity in lateral view, directed posterolaterally.
In dorsal view, shape of petiole scoop with convex side, longer than broad. Anterolateral corner of petiole undeveloped, tapering anteriorly. Petiolar spiracle big, as large as half of propodeal spiracle in diameter. Postpetiole weakly bilobed but without longitudinal median sulcus. Petiole slightly wider than, or as wide as postpetiole in dorsal view.
Integument essentially smooth and shiny. Clypeus generally smooth and shiny, but with feable rugulae. Malar region smooth and shiny without rugulae. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum generally shiny, but with feable rugulae. Dorsal surface of propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed sparse setae. Clypeus without longer setae on anteriormost portion. Anterior clypeal margin with one single longer setae and two pairs of longer setae, mixed with some shorter setae on the side. No erect setae on pronotal shoulder. Posterolateral tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown.

Distribution. This species is distributed in India and Myanmar.
Comments. This species is unique in Asian C. ranavalonae-group in having a slender petiole. This species is similar to C. tumidula, but distinguished in having the petiole without anterolateral corners.
The syntype workers of C. ebenina corax agree well with syntype workers of C. ebenina. Thus I treated C. ebenina corax as a junior synonym of C. ebenina.  General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is less than width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view. Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine undeveloped, but as small tubercule in some workers; length distinctly shorter than propodeal spiracle. Propodeal spiracle large and circular, situated close to propodeal declivity in lateral view, directed laterally.

Crematogaster hashimi
In dorsal view, shape of petiole scoop with convex side, longer than broad. Anterolateral corner of petiole rounded. Petiolar spiracle small, less than half of propodeal spiracle in size. In dorsal view, postpetiole slightly broader than long. Postpetiole weakly bilobed, but without longitudinal median sulcus. Postpetiole as wide as petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron smooth and shiny. Dorsal surface of propodeum smooth and shiny. Lateral propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus, mesosoma and fourth abdominal tergite with short and appressed sparse setae . Clypeus without longer setae on anteriormost portion. Anterior clypeal margin with two or three pairs of longer setae on median portion, with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole without setae. Postpetiole without longer erect setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown. Etymology. The species is dedicated to Dr. Rosli Hashim, University of Malaya, who helped with field surveys in Malaysia.
Distribution. This species is distributed in Thailand and Malaysia (Peninsula). Comments. This species is similar to C. imperfecta, but differs in having circular propodeal spiracles and broader petiole.
This species inhabits well-developed forests, and forage on trees. General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is less than width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view.
In dorsal view, shape of petiole scoop with convex side, longer than broad. Anterolateral corner of petiole developed. Petiolar spiracle small, less than half of propo- deal spiracle in size. In dorsal view, postpetiole slightly broader than long. Postpetiole weakly bilobed, but without longitudinal median sulcus. Postpetiole as wide as petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron smooth and shiny. Dorsal surface of propodeum smooth and shiny. Lateral propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus, mesosoma and fourth abdominal tergite with short and appressed sparse setae. Clypeus with longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two pairs of longer setae on median portion, with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral tubercle with some appressed setae. Ventral surface of petiole without setae. Postpetiole without longer erect setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown. Etymology. The specific name refers to the undeveloped propodeal spines.

Distribution. This species is known from Malaysia (Borneo).
Comments. This species is very similar to C. hashimi, but differs in having elliptical propodeal spiracles and a slender petiole. Fig. 33 Crematogaster tumidula subsp. pia Forel, 1911: 384. Type locality: Malaysia, Berhentian Tingi, Nigri Sembilan, Malacca. Combination in C. (Acrocoelia), and raised to species by Emery, 1922: 151;in C. (Crematogaster)   General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapincal teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is as long as width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view.

Crematogaster pia Forel
Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine long and stout; length longer than spiracle, dorsum as high as anterior propodeum in lateral view; basal width larger than spiracle in diameter. Propodeal spiracle large, situated close to propodeal declivity in lateral view, directed laterally.
In dorsal view, shape of petiole scoop with convex side, as broad as long. Anterolateral corner of petiole angulate. Petiolar spiracle big, as wide as half of propodeal spiracle in diameter, directed laterally. In dorsal view, postpetiole broader than long, bilobed but without longitudinal median sulcus. Postpetiole slightly wider than petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum generally shiny, but with feable rugulae. Dorsal surface of propodeum smooth and shiny; anterior dorsum with feable rugulae.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed sparse setae. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two or three pairs of longer setae on median portion, mixed with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color reddish brown to brownish.

Distribution. This species is distributed in Thailand and Malaysia (Peninsula).
Comments. This species is similar to C. butteli and C. tumidula, but differs from C. butteli in having an anterolaterally developed petiole, and from C. tumidula in having long propodeal spines.
This species inhabits well-developed forests, make a carton nest and forage on trees.
General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapincal teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is as long as width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view.
In dorsal view, shape of petiole scoop with convex side, as long as broad. Petiolar spiracle big, as wide as half of propodeal spiracle in diameter. In dorsal view, postpetiole broader than long, strongly bilobed laterally, but without longitudinal median sulcus. Postpetiole wider than petiole in dorsal view. Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region with feable rugulae. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum generally shiny, but with longitudinal rugulae. Dorsal surface of propodeum smooth and shiny.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed sparse setae. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two pairs of longer setae, mixed with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral tubercle with some decumbent to appressed shorter setae. Ventral surface of petiole with appressed setae. Postpetiole with some shorter setae posteriorly. Fourth abdominal tergite with appressed setae.
Body color brown. Distribution. This species is only known from the type locality in India.
Comments. This species is similar to C. daisyi and C. dalyi, but differs from C. daisyi in having the spiracles on the petiole located on the middle position in lateral view, and from C. dalyi in having long propodeal spines.
Top specimen of three on one pin was designated as lectotype worker here, but it is noted that the body of the middle specimen was lost from the card point. Fig. 35 Crematogaster tumidula Emery, 1900: 689, fig. 8 General description of worker. Head appearing subquadratic in front view. Mandible weakly striate, with four teeth, apical and subapical teeth large, basal two teeth smaller. Scape exceeding posterior corner of head, with appressed setae, each of which is less than width of scape in length. Compound eye large and slightly projecting beyond lateral margin of head in full face view.

Crematogaster tumidula Emery
Ventrolateral katepisternal ridge indistinct posteriorly. Propodeal spine short and stout; length as long as spiracle, directed posteriorly, dorsum as high as anterior propodeum; basal width smaller than spiracles in diameter. Propodeal spiracle large, situated close to propodeal declivity in lateral view, directed laterally or posterolaterally. In dorsal view, shape of petiole scoop with convex side, longer than broad. Anterolateral corner of petiole developed. Petiolar spiracle as large as half of propodeal spiracle in diameter, directed laterally. Postpetiole bilobed but without longitudinal median sulcus. Postpetiole as wide as petiole in dorsal view.
Integument essentially smooth and shiny. Clypeus smooth and shiny without rugulae. Malar region smooth and shiny. Dorsal surface of promesonotum smooth and shiny. Lateral surface of pronotum smooth and shiny. Mesopleuron and lateral propodeum generally smooth, but with feable rugulae. Dorsal surface of propodeum smooth and shiny; anterior dorsum smooth without rugulae.
Erect pilosity almost absent. Dorsum of head, clypeus and mesosoma with short and appressed sparse setae. Clypeus with one pair of longer setae on anteriormost portion, directed medially. Anterior clypeal margin with two pairs of longer setae on median portion, mixed with some shorter setae on side. No erect setae on pronotal shoulder. Posterolateral tubercle with some appressed shorter setae. Ventral surface of petiole with appressed setae. Postpetiole with some appressed setae. Fourth abdominal tergite with appressed setae.
Body color reddish brown. Distribution. This species is known from Malaysia (Borneo) and Indonesia (Kalimantan, Sumatra).
Comments. This species is similar to C. ebenina, but differs in having developed anterolateral corners of the petiole. It is also similar to C. pia, but differs in having short and stout propodeal spines and the basal portion with smooth surface.

Crematogaster vandermeermohri Menozzi
Crematogaster ( Type material. Crematogaster (Paracrema) vandermeermohri: syntype worker from Indonesia, Toetoepan, Sumatra (probably in IEGG, not seen). Comments. I have not been able to examine type-material of C. vandermeermohri. Judging from the original description and figure, this species is close to C. hashimi and C. imperfecta. Menozzi (1930) described the following features: (1) propodeal spines small and blunt, inflated near the base; (2) postpetiole slightly narrower than petiole in dorsal view. The two features are not found in the related species. The postpetiole is as wide as or wider than petiole in C. hashimi and C. imperfecta. or access to museum collections: Daniel Burckhardt (NHMB); Bernhard Merz (MHNG); Maria Tavano and Roberto Poggi (MCSN); Suzanne Ryder and Gavin Broad (BMNH); Dominique Zimmermann (NHMW). Barry Bolton provided a suggestion on the terminology of katepisternum. Additional material was received from Watana Sakchoowong, Weeyawat Jaitrong, Katsuyuki Eguchi. Thanks are also for due to Rosli Hashim and Wattanachai Tasen who provisioned my collecting trips. I would like to thank the ANeT members for encouragement. Thanks are also due to Mark Lorenz (Forte Inc.) for improving the English. This work was supported in part by JSPS KAKENHI (Grant-in-Aid for Scientific Research (C)) Grant Number 26440221.