Hosoishi, S. and K. Ogata. 2016. Systematics and Biogeography Of

Zoological Journal of the Linnean Society, 2016, 176, 547–606. With 11 ﬁgures.

Systematics and biogeography of the ant genus Crematogaster Lund subgenus Orthocrema Santschi in Asia (Hymenoptera: Formicidae)

SHINGO HOSOISHI* and KAZUO OGATA

Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan

Received 26 March 2015; revised 28 June 2015; accepted for publication 8 July 2015

The subgenus Orthocrema of the ant genus Crematogaster is a well deﬁned group and diverse in the tropical Asia. Its systematics has remained poorly understood because of a lack of modern revisionary work. Crematogaster (Orthocrema) is revised for the Asian region, and 27 species including ten new species are recognized. Five species groups: the C. baduvi group (4 spp.); the C. binghamii group (3 spp.); the C. biroi group (10 spp.); the C. moatensis group (1 sp.); the C. quadriruga group (9 spp.) are established based on worker caste morphology. A key to Asian species of the subgenus Orthocrema based on the worker caste is given. Phylogenetic relationships of Asian Orthocrema are analyzed. The analysis revealed that the C. baduvi-, C. binghamii-, and C. biroi groups are monophyletic, and that the allopatric distribution patterns of closely related species imply that Asian Orthocrema is composed of relatively young taxa. There have been at least three west-to-east dispersal events across Wallace’s line in the C. baduvi-, C. quadriruga- and C. biroi groups.

ADDITIONAL KEYWORDS: allopatric distribution – intermediate worker – nesting site – phylogeny – Wallace’s line.

INTRODUCTION more recently described species and changes in the classification of the subgenus (Blaimer, 2012c). The genus Crematogaster is one of the most diverse Orthocrema ants have generally been considered to ant genera, particularly in the tropics (Hölldobler & be ground dwellers, and they have frequently been col- Wilson, 1990). The regional-scale taxonomy and sys- lected by general sampling methods in faunal surveys tematics of the genus, which includes 487 valid species- (e.g., Malsch, Rosciszewski & Maschwitz, 2003; Eguchi level names (Bolton, 2014) in two subgenera (Blaimer, et al., 2005; Pfeiffer et al., 2011). Although some speci- 2012c, d), have received considerable attention in recent mens have been identified to species level based on years (Longino, 2003; Hosoishi & Ogata, 2009b, 2012; distinct morphological characters (e.g., Hosoishi, Yamane Blaimer, 2010, 2012a, 2013). & Ogata, 2010), many specimens have not been iden- The subgenus Orthocrema consists of 142 species tified to species. around the world (Blaimer, 2012c), and of these, 15 Workers of the subgenus are roughly distinguished species and 4 subspecies are presently known from Asia. even in the field, by having a yellow-colored body and In his review of the ants of the former British India, acrobatic style. The subgenus has been characterized Bingham (1903) listed only one species in Orthocrema. based on the following features: two-segmented antennal A key to Orthocrema species from the Indo-Australian club; petiole with subparallel sides; postpetiole without region was provided in Menozzi (1935), but the key median sulcus (Santschi, 1918; Emery, 1922; Wheeler, is of little use now as it has not been updated to include 1922). Subsequently Blaimer (2012c) diagnosed the subgenus based on a variety of character combinations on a global scale. The aim of this study is to revise the *Corresponding author. E-mail: [email protected] Orthocrema species found in Asia.

The Winkler extraction method is generally suit- London, UK; CASC, California Academy of Sciences, able for collecting many of the species in the subge- San Francisco, CA, USA; FRIM, Forest Research In- nus, but a fogging machine is better suited for collecting stitute Malaysia, Kuala Lumpur, Malaysia; HNHM, some of the rarer species in the canopies of tropical Hungarian Natural History Museum, Budapest, rainforests. We also collected several species from dead Hungary; IEBR, Institute of Ecology and Biological Re- twigs or leaves on trees in Asia. As mentioned by sources, Hanoi, Vietnam; IEGG, Istituto di Entomologia Blaimer (2012b) in her revision of Malagasy taxa, the ‘Guido Grandi’, Bologna, Italy; ITLJ, National Insti- nesting site of the subgenus is more extensively dis- tute of Agro-Environmental Sciences, Tsukuba, Japan; tributed than expected. It is not known whether each KUEC, Institute of Tropical Agriculture, Kyushu Uni- taxon in the subgenus shows a particular nesting be- versity, Fukuoka, Japan; MBBJ, Museum Zoologicum haviour. In this study, our collections comprise a wide Bogoriense, Cibinong, Java, Indonesia; MCSN, Museo range of material, covering subterranean and ground- Civico di Storia Naturale ‘Giacomo Doria’, Genoa, Italy; dwelling ants obtained by Winkler extractions and MCZC, Museum of Comparative Zoology, Harvard Uni- general collections, hand-collected arboreal ants, and versity, Cambridge, MA, USA; MHNG, Muséum canopy ants with fogging. We consider that nesting d’Histoire Naturelle, Geneva, Switzerland; MNHA, site information can reveal interesting phylogenetic Museum of Nature and Human Activities, Hyogo, Japan; relationships. MPMP, National Museum of the Philippines, Manila, Generally, workers of the subgenus are monomorphic Philippines; NHMB, Naturhistorisches Museum, Basel, in size, but morphological intermediate workers of Switzerland; NHMW, Naturhistorisches Museum, Wien, queens and ordinary workers have been found in some Austria; SKYC, Seiki Yamane Collections, Kitakyu- species (Heinze et al., 1999; Longino, 2003; Blaimer, shu Museum of Natural History and Human History, 2012b; Peeters et al., 2013). It is reported that the inter- Kitakyushu, Japan; THNHM, Thailand Natural History mediate workers lay unfertilized trophic eggs. In our Museum, Technopolis, Khlong Luang, Pathum Thani, revision of Asian taxa, we found intermediate workers Thailand. In the lists of ‘Material examined’ the records in five species. Phylogenetic relationships reveal the for each species are arranged alphabetically by country. occurrence patterns of the intermediate workers in Asian Nest series samples are represented as colony code, fauna. e. g. ‘SH10-Mal-48’. The goal of the present study is a phylogenetic study Most observations were made under a Leica M205C and taxonomic revision of the subgenus Orthocrema stereomicroscope. Images were created using a Canon in Asia, including keys to the species, distribution maps, EOS 50D with a Canon MP-E 65 mm ×1–5 macro lens, and discussion of biogeographic patterns. We also discuss then processed using CombineZM software. the monophyly and systematic position of the species Measurements were made under the Leica M205C groups based on worker morphology. Nesting behav- stereomicroscope using micrometres. All measure- iour and information of intermediate worker on ments are expressed in millimetres, recorded to the the phylogeny provide insight into the character second decimal place. Abbreviations used for meas- evolution. urements and indices follow Longino (2003) and Hosoishi (2015). The measurements for petiole and postpetiole generally follow Longino (2003) when applicable, except MATERIAL AND METHODS as follows. head width (HW) (Fig. 1); head length (HL) ‘Asia’ in the present paper refers to the Manchurian (Fig. 1); cephalic index (CI): HW/HL × 100; scape length subregion of the Palaearctic Region, Oriental Region, (SL) (Fig. 2); scape index (SI): SL/HW × 100; eye length and Australian region excluding Australia, Papua New (EL) (Fig. 1); pronotal width (PW) (Fig. 3); Weber’s length Guinea, the Solomon Islands, Vanuatu, Fiji and the of the mesosoma (WL) (Fig. 4); propodeal spine length islands in the Pacific Ocean. We generally followed the (PSL) (Fig. 4); petiole length (PtL) (Fig. 5) (slightly modi- geographical terminology of Ward (2001). The term ‘Wal- fied from Longino, 2003: fig. 2); petiole width (PtW) lace’s line’ refers to the original definition of the line (Fig. 6); petiole height (PtH) (Fig. 5) (see Longino, 2003: (Wallace, 1863), that runs east of Bali, Borneo and the fig. 2); postpetiole length (PpL (Fig. 6) (see Longino, Philippines. 2003: fig. 2); postpetiole width (PpW) (Fig. 6); petiole Type specimens were examined and/or deposited in height index (PtHI): PtH/PtL × 100; petiole width index the collections listed below. Codes for public institu- (PtWI): PtW/PtL × 100; postpetiole width index (PpWI): tions generally follow those in Arnett, Samuelson & PpW/PpL × 100; waist index (WI): PpW/PtW × 100. Nishida (1993). The acronyms and abbreviations of the Chaetotaxy is very important for species identifica- institutions/collections cited in the text are: ACEG, Ant tion in some ant genera, such as Lasius (Salata & Collection of Katsuyuki Eguchi, Graduate School of Borowiec, 2011), Formica (Petrov & Collingwood, 1993; Science and Engineering, Tokyo Metropolitan Univer- Fedoseeva, 2011; Korochkina, Konopleva & Zryanina, sity, Tokyo, Japan; BMNH, Natural History Museum, 2014), Polyrhachis (Sorger & Zettel, 2009), Messor

Figures 1–7. 1–6, Measurements of workers. 1, Head width, head length and eye length; 2, Scape length; 3, Pronotal width; 4, Weber’s length and propodeal spine length; 5, Petiole length and petiole height; 6, Petiole width, postpetiole length and postpetiole width. 7, Map of the chaetotaxy of the mesosomal surface of workers of the Asian Orthocrema species showing the positions of the setae, cited from Menozzi (1935).

Table 1. Summary of terms and abbreviations referring to setae important to the systematics of Asian Orthocrema species

Deﬁnition Term or abbreviation Mesosoma Menozzi (1935) ps1PN paired setae on pronotal shoulders A ps2PN paired setae on anteromedian pronotum B psaMN paired setae on anterior mesonotal ridges C pspMN paired setae on posterior mesonotal ridges D psPR paired setae on anterior dorsolateral corners of propodeum E ps1PS paired setae near base of propodeal spines F ps2PS paired setae on dorsal surface of propodeal spines G

(Baroni Urbani, Aktaç & Çamlitepe, 1989) and (Table 2). Thirty-three morphological characters were Technomyrmex (Bolton, 2007), but has not been com- examined from the worker caste and all of the char- monly used in other genera. Different systems of setal acters including a multistate character were treated nomenclature are used in ant taxonomy. The nomen- as unordered; ‘?’ is given for unknown and inappli- clature applies only to the head, mesosoma, and legs cable cases, and the matrix is shown in Table 2. of closely related species or species groups. Conse- For this analysis, 24 Asian Orthocrema species were quently, the nomenclature is typically very limited and selected. not generally applicable on a wide scale. This study Characters of the queens, males and larvae were not did not aim to develop a generally applicable chaetotaxy included as it was not represented in our collections. system for ants; rather, we present a system cover- Buren (1958) stated that the differences in the male ing only the Asian Orthocrema fauna. genitalia are so slight as to be useless for species di- In his review of Asian Crematogaster ants, Menozzi agnosis in the genus Crematogaster. As for larvae, (1935) also introduced a chaetotaxy system for the dorsal Wheeler & Wheeler (1952) examined the former sub- mesosomal surface among members of the subgenus genera, Crematogaster, Orthocrema, Physocrema and Orthocrema. Our collections suggest that workers of Sphaerocrema, but they did not find any characters Asian Orthocrema species generally possess a pattern for diagnosing each subgenus because of intra- and of rows of long erect setae on the body segments (head, interspecific variations. Whereas nest series samples mesosoma, petiole, postpetiole and fourth abdominal in closely related species collected from Southeast Asia tergite), and that chaetotaxy in Asian Orthocrema showed the interspecific differences in the male and species is informative for both species-level identifi- queen castes (Hosoishi, unpublished data). In the present cation and phylogenetic interpretation. We generally study, we examine the workers only, but the morpho- follow Menozzi (1935) and redefine each seta on the logical analysis shown here can contribute to the better mesosoma surface. The chaetotaxy system of Asian understanding of phylogenetic relationships when the Orthocrema species proposed in the present study is male and queen castes are collected in the future. summarized in Figure 7 (Menozzi, 1935: fig. 1) and the Cladistic analyses were performed with the TNT 1.1 nomenclature is given in Table 1. program (Goloboff, Farris & Nixon, 2008) with outgroup As suggested by Sorger & Zettel (2009), although rooting, default consensus options, Tree Bisection and chaetotaxy is important for species identification, count- Reconstruction (TBR) branch swapping, and the default ing setae is often difficult in old or damaged speci- ‘traditional search’ mode. The TNT program accepted mens. We therefore did not rely on only the number only one taxon as an outgroup for the analysis, and of setae to distinguish between species; careful exami- Crematogaster paradoxa was selected for this purpose nation of the specimens in our collection suggests that as molecular data suggest that C. paradoxa is the sister variations in the relative abundance, length and shape group of the remaining Orthocrema clades (Blaimer, of the setae can be useful additional characters for 2012d). The tree search employed a parsimony ratchet phylogenetic analysis. with 10 000 iterations per run. Parsimony analyses were completed under conditions of equal and implied weighting, with k values ranging from 1 to 10. Tree analy- PHYLOGENETIC ANALYSIS ses and graphical manipulations were performed To determine the systematic position of the species with WinClada version 1.00.08 software (Nixon, 2002), groups within the subgenus Orthocrema in Asia, 25 and consensus cladograms generated from equally species, including fauna from New Guinea, were ex- parsimonious trees were generated using the same amined to code the most informative character states program.

Table 2. The subgenus Orthocrema in Asia: dataset for cladistic analysis. The ﬁrst taxon is an outgroup. ‘?’ signiﬁes unknown or inapplicable

Terminal taxa 123456789101112131415161718192021222324252627282930313233

Crematogaster paradoxa 1000000000 0 1 0 ? 110000000000?000000 C. baduvi 0000010000 0 1010000010000010010100

olgclJunlo h ina Society Linnean the of Journal Zoological C. bandarensis 0001100010 1 1010110001000110100110 C. binghamii 0001000010 1 1100110101000111000101 C. biroi 0011100111 1 0010011001111110101110 C. brevispina 0001000110 1 1100110101000111000101 C. brunensis 0000010000 0 1010000010000010010100 C. celebensis 0001100010 1 1010110001000110100110 C. fritzi 0111100111 1 0010011001100110101110

C. longipilosa 0001000110 1 1110110001000111000101 ASIAN OF BIOGEOGRAPHY AND SYSTEMATICS C. luzonensis 0111100111 1 0000011001100110101110 C. macracantha 0000000000 0 1010000010000010010100 C. masukoi 1001100111 1 1001011001001111101110 C. moatensis 0101000110 1 1001100000000010000100 C. myops 0001100010 1 1010110001000110100110 C. ocellata 0001100111 1 1010011001000110101110

2016, , C. osakensis 1001100111 1 0010011001001110101110 C. philippinensis 0001100010 1 1010110001000110100110 C. quadriruga 0001100010 1 0000110001000110100110 176 C. reticulata 1111101111 1 0000011001101110101110

547–606 , C. schimmeri 1111101111 1 0000011001100110101110 C. storki 0000010000 0 1010000010000010010100 C. suehiro 0101100010 1 1000110001000111100110 C. sundalandensis 0001100010 1 1010110001000111100110 C. vieti 0001100111 1 0010011001001110101110 551 552 S. HOSOISHI AND K. OGATA

CHARACTER LIST 11. Pronotum (0) slightly higher than pronotal collar in lateral view (Fig. 18); (1) distinctly higher than 1. Anterior clypeal margin (0) convex in median pronotal collar in lateral view (Fig. 19). portion (Fig. 8); (1) almost straight or weakly 12. Pronotal shoulders (0) with ridge or rugulae lat- concave in median portion (Fig. 9). erally (Fig. 20); (1) without ridge or rugulae lat- 2. Clypeus striated with longitudinal rugulae; rugulae erally (Fig. 21). (0) not extending to posterior clypeal margin; (1) 13. Posterior portions of mesonotal dorsum extending to posterior clypeal margin. (0) not forming short triangular process 3. Clypeus with erect setae; (0) setae not stout and (Fig. 22); (1) forming short triangular process tapering distally; (1) setae stout and not taper- (Fig. 23). ing distally. 14. Mesonotal ridges (0) not extending posteriorly to 4. Scape length in relation to head width (0) longer, propodeal spines (Fig. 24); (1) extending posteri- SI usually > 100; (1) shorter, SI < 100. orly to propodeal spines (Fig. 25). 5. Basal flagellar segment (antennal segment III) (0) 15. Mesonotal ridges (0) developed (Fig. 26); (1) not longer than broad, 1.5 or more × as long as broad developed (Fig. 27). (Fig. 10); (1) as long as broad or broader than long 16. Mesopleuron (0) sculptured; (1) smooth and shining. (Fig. 11). 17. Metanotal groove laterally (0) not margined by 6. Apical two flagellar segments (0) not differentiat- lamellate ridges (Fig. 28); (1) margined by lamellate ed from other flagellar segments in coloration ridges (Fig. 29). (Fig. 12); (1) differentiated from other flagellar seg- 18. Dorsal surface of propodeum (0) generally smooth ments in coloration (Fig. 13). and shining; (1) mat. 7. Dorsal surface of head essentially (0) smooth; (1) 19. Propodeal spines (0) usually developed, longer than sculptured. diameter of propodeal spiracles; (1) undeveloped 8. Gena and surrounding region of antennal sockets (PSL 0) or weakly developed, shorter than diam- (0) smooth; (1) striated with rugulae. eter of propodeal spiracles. 9. Head shape in full-face view, (0) rounded (Fig. 14); 20. Propodeal spines (0) directed posteriorly (Fig. 30); (1) subquadratic (Fig. 15). (1) divergent (Fig. 31). 10. Anterior margin of pronotal collar (0) concave in 21. Propodeal spiracles (0) oval, relatively small, as dorsal view (Fig. 16); (1) almost straight in dorsal large as or slightly larger than mesothoracic spira- view (Fig. 17). cles in diameter (Fig. 32); (1) elliptical, relatively

Figures 8–15. Characters of head. 8, C. baduvi; arrow indicates convex anterior clypeal margin. 9, C. osakensis; arrow indicates weakly concave anterior clypeal margin. 10, C. brunensis; arrow indicates longer antennal segment III. 11, C. osakensis; arrow indicates broad antennal segment III. 12, C. brevispina. 13, C. baduvi. 14, C. baduvi. 15, C. longippilosa.

Figures 16–23. Characters of mesosoma. 16, C. quadriruga; arrow indicates concave anterior margin of pronotal collar. 17, C. reticulata; arrow indicates straight anterior margin of pronotal collar. 18, C. brunensis; arrow indicates slightly higher pronotum. 19, C. osakensis; arrow indicates distinctly higher pronotum. 20, C. binghamii. 21, C. biroi. 22, C. quadriruga; arrow indicates posterior portion of mesonotal dorsum. 23, C. brevispina; arrow indicates short triangular process on posterior portion of mesonotal dorsum.

17:0 17:1

20:0 20:1

Figures 24–31. Characters of mesosoma. 24, C. reticulata; arrow indicates mesonotal ridge. 25, C. baduvi; arrow indicates extending mesonotal ridge. 26, C. brunensis. 27, C. masukoi. 28, C. baduvi. 29, C. fritzi. 30, C. longipilosa; arrow indicates propodeal spine directed posteriorly. 31, C. brunensis; arrow indicates propodeal spines divergent.

23:0

21:0 21:1

23:1 29:1 29:0

30:0 30:1 24:0 24:1 26:1 26:0

Figures 32–39. Characters of mesosoma, petiole, and postpetiole. 32, C. storki; arrow indicates oval and small propodeal spiracle. 33, C. osakensis; arrow indicates elliptical and large propodeal spiracle. 34, C. longipilosa; arrow indicates petiole with anterolateral corner round. 35, C. biroi; arrow indicates petiole with anterolateral corner angulate. 36, C. brunensis; arrow indicates undeveloped subpetiolar process (24:0), and undeveloped subpostpetiolar process (30:0). 37, C. luzonensis; arrow indicates developed subpetiolar process (24:1), and developed subpostpetiolar process (30:1). 38, C. paradoxa; arrow indicates posterior portion of petiole posterior raised to form small node. 39, C. osakensis; arrow indicates ﬂat posterior portion of petiole.

large, more than 2 × as large as mesothoracic spira- 27. Posterior portion of petiole (0) with short process; cles in diameter (Fig. 33). short process higher than posterior end (Fig. 40); 22. Erect setae on mesosoma; (0) tapering distally; (1) (1) without distinct short process; highest at pos- stout, not tapering distally. terior end (Fig. 41). The short process is widely 23. Petiole with (0) anterolateral corners round found in Asian Orthocrema species, but the de- (Fig. 34); (1) anterolateral corners angulate velopment is variable in some species. (Fig. 35). 28. Postpetiole (0) not bilobed posteriorly (Fig. 42); (1) 24. Subpetiolar process (0) undeveloped (Fig. 36); weakly bilobed posteriorly, but without longitudi- (1) developed, distinctly produced downward nal median sulcus (Fig. 43). (Fig. 37). 29. Postpetiole in lateral view (0) dorsally not 25. Petiole (0) relatively long and slender (PtWI strongly convex, only as high as petiole (Fig. 37); usually < 80); (1) relatively short and broad (PtWI (1) strongly convex, distinctly higher than petiole usually > 80). (Fig. 36). 26. Posterior portion of petiole (0) posteriorly raised 30. Subpostpetiolar process (0) not developed or to form small node (Fig. 38); (1) completely ﬂat venter convex (Fig. 36); (1) developed as process (Fig. 39). (Fig. 37).

27:0 27:1

28:0

28:1 31:0 31:1

Figures 40–45. Characters of petiole and postpetiole. 40, C. storki; arrow indicates short process on posterior portion of petiole. 41, C. masukoi; arrow indicates undeveloped process on posterior portion of petiole. 42, C. binghamii; arrow indicates postpetiole not bilobed posteriorly. 43, C. bandarensis; arrow indicates postpetiole bilobed posteriorly. 44, C. paradoxa; arrow indicates ﬂat subpostpetiolar portion. 45, C. baduvi; arrow indicates convex subpostpetiolar portion.

31. Subpostpetiolar portion; (0) venter flat in lateral each other, all of the topologies obtained by implied view (Fig. 44); (1) venter convex or developed as weighting recovered the C. quadriruga group as process in lateral view (Fig. 45). paraphyletic, an unresolved polytomy sister to the C. 32. Fourth abdominal tergite with (0) erect or suberect biroi group. setae only; (1) erect or suberect setae mixed with In the equal- and implied-weighting parsimony analy- decumbent or appressed setae. ses, the clade for the C. baduvi group was character- 33. Standing pilosity on body surface (0) sparse; (1) ized by the following character states: mesopleuron abundant. sculptured (character 16: 0), propodeal spines directed laterally (character 20: 1); postpetiolar dorsum highly convex (character 29: 1); the clade for the C. binghamii group by the following character states: posterior portion RESULTS of mesonotal dorsum forming a short triangle-shaped The equal-weighting analysis recovered four equally process (character 13: 1); standing pilosity long and parsimonious cladograms and the strict consensus abundant (character 33: 1); and the clade for the C. cladogram collapsed seven nodes (Fig. 46) with a total biroi group by the following character states: anteri- number of 55 steps, a consistency index of 60 and a or margin of pronotal collar almost straight (charac- retention index of 85. The inferred species groups con- ter 10: 1), mesopleuron sculptured (character 16: 0), sisted of the same species in all trees. dorsal surface of propodeum striated with rugulae (char- The implied-weighting cladograms with values of acter 18: 1), subpostpetiolar process developed as process k = 10 produced similar topologies (Fig. 47), with a total (character 30: 1). The clade composed of the C. number of 53 steps, a consistency index of 62 and a quadriruga group and C. biroi group was supported retention index of 87. Although the general topology by the following set of synapomorphies: basal flagel- was consistent among all of the resulting trees, the lar segment (antennal segment III) broader or as long cladistic relationships varied within the species groups. as broad (character 5: 1), weakly bilobed postpetiole Although polytomy was obtained in the strict con- (character 28: 1), erect setae mixed with appressed setae sensus tree (Fig. 46), this tree did not differ from the on fourth abdominal tergite (character 32: 1). equal-weighting tree in terms of the relationships The C. baduvi group was consistently placed at a between the species groups. basal position to the rest of the taxa. All analyses re- The three species groups defined here, i.e., C. baduvi covered Crematogaster moatensis as the sister taxon group; C. binghamii group; and C. biroi group, were to the clade composed of [C. binghamii group + (C. all monophyletic. Although they differed slightly from quadriruga group + C. biroi group)].

Figure 46. Strict consensus tree of four equally parsimonious cladograms obtained by equal-weighting analysis (length: 55 steps; consistency index 60; retention index: 85). Closed circles represent unique apomorphies and open circles homoplasies. Character numbers are given above each circle.

DISCUSSION differ significantly from the implied-weighting trees. PHYLOGENETIC RELATIONSHIPS WITHIN THE As our primary goal was to clarify the monophyly of the species groups defined, slight variations in tree to- ASIAN Orthocrema pology were not considered important. In this study, the strict consensus tree obtained by the Blaimer (2012d) used five nuclear genes (3384 bp) to equal-weighting analysis was used to infer the estimate the molecular phylogeny of 124 Crematogaster phylogenetic relationships among the Asian Orthocrema taxa from around the world. Taxa were selected to cover species as the topology of the consensus tree did not a broad geographic range in order to clarify the

Figure 47. Strict consensus tree of k = 10 obtained by implied-weighting analysis (length: 53 steps; consistency index 62; retention index: 87). Closed circles represent unique apomorphies and open circles homoplasies. Character numbers are given above each circle. phylogenetic relationship among species as accurately extent (Blaimer, 2012d, Fig. 2). Crematogaster reticulata as possible; her analysis included eight Asian Orthocrema and C. osakensis were placed within the same clade taxa. Although the cladistic analysis presented here and C. binghamii was a sister to C. longipilosa. However, based on worker morphology is limited to Asian fauna, C. fritzi did not form a clade with C. baduvi in our it corresponds with her phylogenetic analysis to some cladistic analysis. These ﬁndings implied that minor

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 559 topological differences might arise between morpho- did not share any apomorphies with the other species logical and molecular analyses at higher taxonomic levels. but its position was consistent, we place C. moatensis Such topological differences might be due to relatively in the C. moatensis group, which is represented by a fewer characters based on worker caste only, and it is single species. Further studies on closely related taxa also needed to examine queens and males. Whereas would clarify the systematic position of the C. moatensis Ward (2007) suggests that it is challenging to infer group. phylogenetic relationships from morphological char- Although we do not estimate the relative diver- acters alone due to convergence. Rather, molecular analy- gence time by using morphological differences, their sis using multiple nuclear genes resolve the differences restricted distribution ranges of C. baduvi group more clearly (Blaimer, 2012d), but it is beyond the scope (Sundaic region) and C. moatensis group (Sulawesi) and of this study to use molecular data due to a lack of a few derived character states imply that the two species fresh material. However this study will facilitate future groups are older taxa than the other three species studies in Asian Orthocrema species, and will provide groups (C. binghamii, C. biroi and C. quadriruga a starting point for molecular studies to test the groups). monophyly of the species groups defined here and the phylogenetic relationships. The species groups were defined based on putative BIOGEOGRAPHY AND NATURAL HISTORY apomorphies and discrete morphological boundaries. We defined the species groups by distinct character The distributions of individual species of Asian states even if cladistic analyses did not find any Orthocrema species imply several different biogeo- synapomorphic characters. We established the C. graphic patterns (Figs 49–52). Closely related species quadriruga group as having the following character appear to be allopatric; in implied-weighted trees states: long scape (SI 81–100); basal flagellar segment (Fig. 47), there are four pairs of sister species, and three (antennal segment III) broader than long; subpetiolar of them clearly exhibit allopatric ranges: C. binghamii process developed; subpostpetiolar portion convex in (Indochina) and C. brevispina (Philippines); C. suehiro lateral view. Although these character states can be (Japan) and C. sundalandensis (Borneo, Sumatra); C. used to separate the species group from other Asian reticulata (W. Malaysia, Borneo, Sumatra) and C. Orthocrema species, our cladistic analysis did not iden- schimmeri (Taiwan). Ward (2001) suggested that the tify any autapomorphies in the C. quadriruga group. Tetraponera species of Indo-Australian region are rela- Despite a lack of synapomorphy, we treat the C. tively old taxa based on sympatric distributions of closely quadriruga group as a distinct species group, al- related taxa. However, in contrast with Tetraponera, though the results were based entirely on morpho- our observations suggest that the Asian Orthocrema logical analysis inferred by relatively few data. Future contains relatively young taxa. Recent molecular analy- studies employing more comprehensive morphologi- ses inferred the divergence time of various ant taxa cal or molecular data therefore need to be undertak- to be as follows: Crematogaster: 34.7 Mya, with 95% en in order to reconstruct the phylogenetic relationships highest posterior density of 28.1–42.4 Mya (Ward et al., between the members of the C. quadriruga group and 2015); Orthocrema clade: 22.3 to 30.2 Mya (Blaimer, other species groups. In addition, such groupings are 2012d); Pseudomyrmecinae: 50.5 to 59.2 Mya (Moreau taxonomically convenient for practical use. et al., 2006); Tetraponera node: 54 Mya (Ward & Brady, Our cladistic analysis indicated that C. moatensis 2003). These estimates support the hypothesis that the did not form a monophyletic clade with any other Asian Orthocrema fauna appeared relatively recently. species. The difficulty associated with establishing an As in the studies on Tetraponera by Ward (2001), accurate position for C. moatensis was due to the char- the species richness of Asian Orthocrema is highest acter states observed in this species. The clade com- in Borneo (10 spp.), followed by Sumatra (7 spp.) and posed of C. moatensis and the C. binghamii, C. Peninsular Malaysia (6 spp.); indeed, it appears that quadriruga and C. biroi groups was supported by the the Sundaic region is the center of diversity for Asian following synapomorphies: relatively short scape length; Orthocrema species (Table 3). two apical flagellar segments non-differentiated in col- The material examined in this study was mainly com- oration; square head shape; pronotum distinctly higher posed of species from the Oriental biogeographic region, than pronotal collar. Whereas C. moatensis could be and relatively few species from the Australian biogeo- separated from the clade for the C. binghamii, C. graphic region were considered. Estimates of quadriruga and C. biroi groups by the following char- phylogenetic relationships imply that the distribu- acter states (non-apomorphies): metanotal groove not tions of C. storki, C. celebensis and C. fritzi traverse covered by lamellate ridges; oval propodeal spiracles; across Wallace’s line, implying that there have been long and slender petiole, therefore C. moatensis was at least three dispersal events to the Australian region sister to the three species groups. Since C. moatensis by the C. baduvi, C. quadriruga and C. biroi groups

Table 3. Distribution of Orthocrema species in the Asian region

Indochina, excluding Indian Japan, Peninsular Peninsular Java,

© Species group Species subcontinent Taiwan Malaysia Malaysia Sumatra Bali Borneo Philippines Sulawesi 06TeLnenSceyo London, of Society Linnean The 2016 baduvi baduvi xxxx brunensis x macracantha x storki x binghamii binghamii xx brevispina x longipilosa xx x biroi biroi x fritzi xx x x luzonensis x masukoi x

olgclJunlo h ina Society Linnean the of Journal Zoological ocellata x osakensis x reticulata xx schimmeri x udo x vieti x quadriruga bandarensis xxxx celebensis x gavapiga x javanica x myops xx x philippinensis x quadriruga xx suehiro x

2016, , sundalandensis xx moatensis moatensis x

176 Total number of species 2 3 4 6 7 5 10 3 4 547–606 , SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 561

Figure 48. Distribution of biogeographic regions, nesting sites and intermediate workers in Asian Orthocrema species. Abbreviations: AUS, Australian; ORI, Oriental; PAL, Palearctic. *Peeters et al. (2013).

(Fig. 48). As Briggs (1987) suggested, it seems that the lection is not ﬁxed in each species group, and that vari- west-to-east dispersal across Wallace’s line is relative- ations occur within species groups (Fig. 48). ly widespread in Asian Orthocrema species. Hosoishi et al. (2010) revised the subterranean species The subgenus Orthocrema is generally considered to from the Asian Orthocrema fauna. Crematogaster contain ground dwellers that nest in soil, but some masukoi and C. myops have reduced compound eyes species are arboreal and nest in dead twigs on trees (approximately 6–10 ommatidia) and depigmented yel- (Blaimer, 2012b). Although information on nesting lowish bodies. The two species are similar in that they biology of Asian Orthocrema species is limited, our col- have reduced eyes, but they are quite different from lections show that a variety of nesting sites is used. each other in other morphological aspects (Hosoishi et al., Crematogaster baduvi and C. storki were collected by 2010). Our cladistic analysis showed that C. masukoi using fogging machine, and one nest of C. baduvi was belongs to the C. biroi group, whereas C. myops belongs found in decayed wood (Yamane, 2013); these species to the C. quadriruga group, suggesting that their sub- are considered to nest in dead twigs or in leaf litter terranean mode of life evolved independently and is mats on trees. Conversely, C. longipilosa, C. bandarensis, an example of convergence (Fig. 48). C. biroi and C. osakensis are all typical ground dwell- Most Orthocrema species are supposed to be general ers that nest in soil on the ground or in leaf litter. predators. Interestingly, in the subterranean species Crematogaster quadriruga, C. suehiro, C. fritzi, C. (C. masukoi and C. myops), their mandibles show dif- reticulata all nest in dead twigs or in dead leaves on ferent character states from other species. The basal trees. Cladistic analysis also showed that nest site se- tooth of the mandibles is arranged away from the third

Figure 49. Geographical distribution of the species of the C. baduvi group and C. moatensis. apical one in those species, although their teeth on mas- cialized trophic function (Peeters et al., 2013). In the ticatory margin are arranged at an equal distance in Neotropical species, C. smithi, many of the unfertilized most species. Little information is known about the eggs are eaten by the queen and larvae, but some eggs biology of those species, but the unique arrangement can develop into males (Heinze et al., 2000; Oettler, of the mandibles may be related to their predatory be- Dijkstra & Heinze, 2013). In this revision of the Asian haviours (Bolton, 1988). taxa, we found intermediate workers in ﬁve species, Workers in the subgenus Orthocrema are generally C. biroi, C. reticulata, C. schimmeri (See Peeters et al., monomorphic in size, but intermediate workers have 2013), C. vieti and C. quadriruga. The intermediate been found in some species in North America (Heinze workers are clearly separated from ordinary workers et al., 1999), Madagascar (Blaimer, 2012b), Brazil by the following features: larger body size, developed (Longino, 2003; Quinet et al., 2009) and Taiwan (Peeters ocelli (sometimes vestigial), highly convex mesonotum. et al., 2013). These workers are intermediate in size The former four belong to the C. biroi group, while between workers and queens. It is reported that the the last belongs to the C. quadriruga group, suggest- intermediate workers laid unfertilized trophic eggs for ing that intermediate workers are widely scattered larvae (Heinze et al., 1999; Peeters et al., 2013). His- across Asian Orthocrema species (Fig. 48). tological studies reveal that the intermediate workers The subgenus Orthocrema is abundant among Asian have developed ovaries but lack a spermatheca, it is ground ant fauna in the temperate to tropical regions. suggesting that they are a soldier caste with a spe- Crematogaster osakensis was one of the most

Figure 50. Geographical distribution of the species of the C. binghamii group. abundant species collected by using Winkler extrac- purpose of taxonomic convenience, we present the fol- tions in grasslands of temperate regions, Japan (Hosoishi lowing characteristics for Asian fauna: et al., 2015). Several colonies of C. vieti and C. bandarensis were frequently collected as nest series 1. Mandibles with four teeth on the masticatory in northern Vietnam (Eguchi et al., 2005) and Bogor, margin; the teeth located at equal distance each Indonesia (Ito et al., 2001), respectively. other in most species, but basal tooth located apart from other teeth in subterranean species (Hosoishi et al., 2010). SYSTEMATICS 2. Palp formula 5, 3 (Bolton, 2003). 3. Anterolateral margins of clypeus not protruded an- SYNOPSIS OF ASIAN SPECIES OF THE SUBGENUS teriorly (Hosoishi & Ogata, 2012). Orthocrema 4. Clypeus usually striated with longitudinal rugulae; The worker caste of the subgenus Orthocrema was di- the length of rugulae variable in different species. agnosed by Blaimer (2012c) on a global scale. For the 5. Antenna 11-segmented.

Figure 51. Geographical distribution of the species of the C. biroi group.

6. Antennal club 2-segmented. Apical two flagellar 12. Ventrolateral katepisternal ridge distinct (Hosoishi, segments clearly distinguished in coloration from 2015). other flagellar segments especially in the C. baduvi 13. Propodeal spines usually developed, but undevel- group. oped or small tubercles in C. binghamii. 7. Sensilla basiconica and sensilla trichodea curvata 14. Petiole usually with subparallel sides. distributed in the apical two flagellar segments 15. Petiole usually with short process posteriorly. (Hosoishi & Ogata, 2009c). 16. Postpetiole not bilobed. If weakly bilobed, usually 8. Compound eye generally distinct and large, but without distinct longitudinal median sulcus. strongly reduced in subterranean species (Hosoishi 17. Fourth abdominal tergite with erect to suberect et al., 2010). setae. 9. Ocelli rarely present in intermediate workers. 18. Body principally yellow, but sometimes partly brown 10. Occipital carina distinct. to black in C. binghami group, C. biroi group, C. 11. Mesonotal ridges developed dorsolaterally, but moatensis group and C. quadriruga group, but weakly developed in C. masukoi and C. moatensis. brown in C. baduvi group.

Figure 52. Geographical distribution of the species of the C. quadriruga group.

19. Monomorphic in size; intermediate workers occa- monophyletic, but also considered as useful units for sionally present. taxonomic purposes in the case of hyperdiverse genera. 20. Nesting in soil; some species nesting in dead twigs on lower vegetation. Crematogaster baduvi group C. baduvi Forel, 1912b. C. brunensis sp. nov. SYNONYMIC LIST OF ASIAN Orthocrema SPECIES C. macracantha Creighton, 1945. In this study, five species groups are established within C. storki sp. nov. the Asian Orthocrema. The species groups were defined based on putative apomorphies and discrete morpho- Crematogaster binghamii group logical boundaries. The species groups recognized here C. binghamii Forel, 1904. will provide a possible basis for new monophyletic taxa. C. brevispina sp. nov. The C. baduvi, C. binghamii and C. biroi groups are C. longipilosa Forel, 1907. relatively easily defined, but the C. quadriruga group is probably not monophyletic as currently defined. The Crematogaster biroi group C. moatensis group was represented by a single species C. biroi Mayr, 1897. since the position of C. moatensis in cladograms was = C. aitkenii Forel, 1902a. syn. nov. always consistent. The taxa should ideally be = C. biroi smythiesii Forel, 1902a. syn. nov.

= C. urvijae Bharti, 2003. syn. nov. Anterior margin of pronotal collar almost straight in C. fritzi Emery, 1901. dorsal view. Dorsal surface of propodeum striated with C. luzonensis sp. nov. rugulae. Mesosoma sculptured. Subpetiolar process de- C. masukoi Hosoishi, Yamane & Ogata, 2010. veloped acutely. Subpostpetiolar process developed C. ocellata sp. nov. acutely. C. osakensis Forel, 1900. This species group is easily distinguished by the C. reticulata Hosoishi, 2009. subpostpetiolar process developed acutely and sculp- C. schimmeri Forel, 1912a. tured surface of mesosoma from other Asian Orthocrema C. udo Forel, 1905. species. C. vieti sp. nov. Crematogaster moatensis group Crematogaster moatensis group Two apical flagellar segments not differentiated in C. moatensis sp. nov. coloration. Pronotum distinctly higher than pronotal collar. Metanotal groove not covered by lamellate Crematogaster quadriruga group ridges. Propodeal spiracles oval. Petiole long and C. bandarensis Forel, 1913. stat. nov. slender. = C. biroi andelis Santschi, 1928. syn. nov. This species group is unique in having the metanotal C. celebensis sp. nov. groove not covered by lamellate ridges, oval-shaped C. gavapiga Menozzi, 1935. propodeal spiracles and long and slender petiole from C. javanica Menozzi, 1935. other Asian Orthocrema fauna. C. myops Forel, 1911a. C. philippinensis sp. nov. Crematogaster quadriruga group C. quadriruga Forel, 1911c. stat. nov. Relatively long scape (SI 81–100). Basal flagellar C. suehiro Terayama, 1999. segment (antennal segment III) broader than long. = C. miroku Terayama, 2013. syn. nov. Subpetiolar process developed. Subpostpetiolar portion C. sundalandensis sp. nov. wholly convex in lateral view. Mesosoma generally smooth and shining. The species groups of Asian Orthocrema species This species group is distinguished by the Apomorphies of the species groups obtained by cladistic subpostpetiolar portion wholly convex and smooth and analysis are given in italics. shining surface of mesosoma from other Asian Orthocrema species. Crematogaster baduvi group Relatively long scape (SI 98–118). Basal flagellar segment (antennal segment III) longer than broad. SPECIES ACCOUNTS Propodeal spines long and directed laterally. Postpetiolar CREMATOGASTER BADUVI FOREL, 1912b dorsum highly convex in lateral view; postpetiole dis- (FIG. 96) tinctly higher than petiole in lateral view. This species group is easily distinguished by the Crematogaster baduvi Forel, 1912b: 106; one syntype propodeal spines directed laterally and highly convex worker, Nusa Kambangan, Java, Indonesia (MHNG, postpetiolar dorsum from other Asian Orthocrema examined). species. Crematogaster baduvi; Forel, 1913: 77 [Description of queen]. Crematogaster binghamii group Crematogaster baduvi [sic]; Crawley, 1924: 394. Basal flagellar segment (antennal segment III) longer Crematogaster baduvi; Menozzi, 1935: 104 [Descrip- than broad. Posterior margins of mesonotum forming tion of male]. short triangle-shaped process in lateral view. Propodeal Crematogaster baduvi; Emery, 1922: 131 [Combina- spines undeveloped, or developed and directed poste- tion in C.(Orthocrema)]. riorly. Standing pilosity abundant on body surface. Crematogaster baduvi; Blaimer, 2012c: 55 [Combi- This species group is easily distinguished by the short nation in C.(Orthocrema)]. triangle-shaped process on posterior margins of mesonotum and long and abundant standing pilosity Worker measurements (n = 17): HW 0.47–0.57; HL 0.48– from other Asian Orthocrema species. 0.59; CI 92–100; SL 0.47–0.58; SI 98–108; EL 0.11– 0.14; PW 0.26–0.35; WL 0.60–0.72; PSL 0.15–0.21; PtL Crematogaster biroi group 0.23–0.28; PtW 0.15–0.20; PtH 0.13–0.16; PpL 0.13– Relatively short scape (SI 70–89). Basal flagellar 0.16; PpW 0.16–0.20; PtHI 54–67; PtWI 65–83; PpWI segment (antennal segment III) broader than long. 113–146; WI 90–107.

KEY TO SPECIES BASED ON THE WORKER CASTE (EXCEPT FOR C. GAVAPIGA MENOZZI, C. JAVANICA MENOZZI, AND C. UDO FOREL THAT WERE NOT EXAMINED) 1. Propodeal spines directed laterally (Fig. 31). Postpetiolar dorsum highly convex in lateral view; postpetiole distinctly higher than petiole in lateral view (Fig. 36) (C. baduvi group) ...... 2 – Propodeal spines directed posteriorly (Fig. 30). Postpetiolar dorsum not highly convex in lateral view; postpetiole as high as petiole in lateral view (Fig. 37)...... 5 2. Transverse distance between tips of propodeal spines distinctly longer than head width in dorsal view (Fig. 53). Mesosoma densely sculptured (Borneo) ...... C. macracantha Creighton – Transverse distance between tips of propodeal spines shorter than head width in dorsal view (Fig. 54). Mesosoma generally smooth and shining ...... 3 3. Propodeal spines curved upward at tip (Fig. 55) (Sulawesi) ...... C. storki, sp. nov. – Propodeal spines not curved upward at tip (Fig. 56)...... 4 4. Head smaller (HW 0.43–0.47; HL 0.46–0.49). Scape relatively long (SI 113–118). Propodeal dorsum smooth. Propodeal spines long (PSL 0.20–0.22). Lateral surface of petiole weakly sculptured (Borneo)...... C. brunensis, sp. nov. – Head larger (HW 0.47–0.57; HL 0.48–0.59). Scape relatively short (SI 98–108). Propodeal dorsum striated with feeble rugulae. Propodeal spines relatively shorter (PSL 0.15–0.21). Lateral surface of petiole sculptured (W. Ma- laysia, Borneo, Java, Sumatra) ...... C. baduvi Forel 5. Metanotal groove not covered by lamellate ridges dorsolaterally (Fig. 28) (Sulawesi) ...... C. moatensis, sp. nov. – Metanotal groove covered by lamellate ridges dorsolaterally (Fig. 29)...... 6 6. Posterior portions of mesonotal dorsum forming short triangle-shaped process (Fig. 23). Standing pilosity long and abundant on head, mesosoma, petiole, postpetiole and remains of the gaster (Fig. 57) (C. binghamii group)...... 7 – Posterior portions of mesonotal dorsum not forming short triangle-shaped process (Fig. 22). Standing pilosity short and sparse (Fig. 58)...... 9 7. Propodeal spines developed, distinctly longer than diameter of propodeal spiracles (Fig. 59) (S. Thailand, W. Ma- laysia, Borneo, Java, Sumatra)...... C. longipilosa Forel – Propodeal spines undeveloped, or weakly developed, as long as or shorter than diameter of propodeal spiracles (Figs 60, 61)...... 8 8. Propodeal spines undeveloped or weakly developed, shorter than diameter of propodeal spiracles (PSL 0–0.03) (Fig. 60) (Nepal, India, Bangladesh, Thailand, Vietnam) ...... C. binghamii Forel – Propodeal spines slightly more developed, as long as diameter of propodeal spiracles (PSL 0.04–0.07) (Fig. 61) (Philippines)...... C. brevispina, sp. nov. 9. Relatively short scape (SI 70–91). Mesosoma sculptured (Fig. 62). Subpetiolar process developed acutely (Fig. 64). Subpostpetiolar process developed acutely (Fig. 64) (C. biroi group)...... 10 – Relatively long scape (SI 81–100). Mesosoma essentially smooth and shining (Fig. 63). Subpetiolar process developed (Fig. 65). Subpostpetiolar process not developed as process; the venter convex (Fig. 65) (C. quadriruga group)...... 18 10. Compound eyes reduced, with less than 15 ommatidia (Figs 66, 67)...... 11 – Compound eyes not reduced, with more than 20 ommatidia (Fig. 68)...... 12 11. Number of ommatidia approximately 5–6 (Fig. 66) (Borneo)...... C. masukoi Hosoishi, Yaname & Ogata – Number of ommatidia approximately 12–15 (Fig. 67) (N. Vietnam)...... C. ocellata, sp. nov. 12. Dorsal surface of head sculptured (Fig. 69) ...... 13 – Dorsal surface of head generally smooth and shining (Fig. 70) ...... 14 13. Head, mesosoma, petiole and postpetiole strongly sculptured reticulately. Propodeal spiracles apart from metapleural gland bulla (Fig. 71). Petiole squared, but without angulate anterolateral corners (Fig. 73) (Thailand, W. Malay- sia, Borneo)...... C. reticulata Hosoishi – Head, mesosoma, petiole and postpetiole weakly sculptured reticulately. Propodeal spiracles touching metapleural gland bulla (as in Figure 72). Petiole squared with angulate anterolateral corners (Fig. 74) (Taiwan)...... C. schimmeri Forel 14. Dorsal surface of head generally smooth, but feebly sculptured at surrounding portion of compound eyes (Fig. 75). Area in front of occipital carinae sculptured (Fig. 77) (India, Sri Lanka)...... C. biroi Mayr – Dorsal surface of head smooth, without sculptured area in front of occipital carinae (Fig. 76)...... 15 15. Head, mesosoma, petiole and postpetiole with erect setae: setae tapering distally (Fig. 78)...... 16 – Head, mesosoma, petiole and postpetiole with erect setae; setae stout and not tapering distally (Fig. 79) ...... 17 16. Propodeal spines slender (Fig. 80). Petiole tapering posteriorly in dorsal view (Fig. 82). Subpostpetiolar process angulate (Fig. 80) (China, Korea, Japan)...... C. osakensis Forel

– Propodeal spines thick (Fig. 81). Petiole with subparallel sides in dorsal view (Fig. 83). Subpostpetiolar process developed acutely (Fig. 81) (Vietnam)...... C. vieti, sp. nov. 17. Scape with suberect to decumbent setae (Fig. 84). Erect setae on anterior mesonotal ridge as long as on posterior mesonotal ridge (Fig. 86). Fourth abdominal tergite with erect setae abundantly (c. > 20) (Philippines)...... C. luzonensis, sp. nov. – Scape with appressed setae (Fig. 85). Erect setae on anterior mesonotal ridge longer than on posterior mesonotal ridge (Fig. 87). Fourth abdominal tergite with erect setae sparsely (c. 4–6) (W. Malaysia, Borneo, Sumatra, Sulawesi)...... C. fritzi Emery 18. Compound eyes reduced, with c. 6 ommatidia (Fig. 88) (W. Malaysia, Borneo, Sumatra)...... C. myops Forel – Compound eyes not reduced, with more than 20 ommatidia (Fig. 89)...... 19 19. Propodeal spiracles large and touching metapleural gland bulla (Fig. 90) ...... 20. – Propodeal spiracles relatively small and apart from metapleural gland bulla (Fig. 91)...... 23 20. Lamellate ridge superficially covering metanotal groove in some specimens, but metanotal groove deep, U-shaped in lateral view (Fig. 92). Propodeal spines long and stout (Fig. 92) (PSL 0.09–0.1) (Sulawesi)...... C. celebensis, sp. nov. – Lamellate ridge superficially covering metanotal groove in some specimens, but metanotal groove not so deep, V-shaped in lateral view (Fig. 93). Propodeal spines relatively short to long (Fig. 93) (PSL 0.05–0.11) ...... 21 21. Propodeal spines relatively short (PSL 0.05–0.08) and stout (Fig. 90) (S. Thailand, W. Malaysia, Borneo, Java, Sumatra, Krakatau)...... C. bandarensis Forel – Propodeal spines relatively long (PSL 0.08–0.11) and slender (Fig. 93)...... 22 22. Scape with suberect setae only. Body color yellow (Philippines)...... C. philippinensis, sp. nov. – Scape with suberect setae mixed with two to three longer setae. Body bicolored, with head and gaster brown, and mesosoma, petiole and postpetiole yelow (Borneo, Sumatra) ...... C. sundalandensis, sp. nov. 23. Longitudinal rugulae on clypeus not reaching area between frontal carinae. Body bicolored, with head, mesosoma, petiole, postpetiole and first gastral tergite yellow, and remains of gaster brown in typical form, but rarely whole body yellowish. Petiole with subparallel sides in dorsal view (Fig. 94) (Laos, Cambodia, Thailand, Sumatra)...... C. quadriruga, stat. nov. – Longitudinal rugulae on clypeus extending to posterior clypeal margin. Body color yellow. Petiole tapering anteriorly in dorsal view (Fig. 95) (Japan)...... C. suehiro Terayama

General description of worker: Workers monomorphic. disc in lateral view. Subpetiolar process weakly de- Head round in full-face view. Mandibles with four veloped as angulate tubercle. Postpetiole in lateral view teeth arranged at an equal distance, apical and with strongly convex dorsum, distinctly higher than subapical teeth large, basal two teeth smaller. Ante- petiole, in dorsal view as wide as petiole, globular, not rior clypeal margin convex in medial portion. Com- bilobed. Venter of postpetiole convex but without dis- pound eyes distinctly projecting beyond lateral margins tinct process. of head in full-face view. Scapes exceeding posterolateral Integument essentially smooth and shining. Dorsal corners of head. surface of head smooth and shining. Mandibles Pronotal collar with concave anterior margin in dorsal with feeble rugulae and smooth interspaces. Clypeus view, slightly lower than pronotum in lateral view. smooth and shining without distinct longitudinal Pronotal dorsum without distinct ridges laterally. rugulae, but short rugulae present anteriorly Mesonotal dorsum with lateral ridges that extend in some specimens. Dorsal and lateral surfaces of posteriad to tips of propodeal spines; anterior ridges pronotum smooth and shining; anterolateral shoul- usually as high as posterior ridges. Pronotum and ders of pronotum with rugulae. Mesopleura general- mesonotum in lateral view forming evenly arched, ly sculptured, but sometimes smooth except for their continuous dorsal outline. Metanotal groove in dorsal marginal areas in some specimens. Dorsal surface of view transverse, almost straight in median portion, propodeum generally smooth and shining, but with forming shallow concavity that is laterally margined rugulae dorsolateral areas. Dorsal surface of petiole by ridges. Propodeal spiracles oval, situated at smooth and shining. Lateral surface of petiole sculp- posterolateral corners of propodeum, apart from tured. Dorsal and lateral surfaces of postpetiole smooth metapleural gland bullae. Propodeal spines devel- and shining. oped, longer than diameter of propodeal spiracles, in Standing pilosity sparse. Dorsal face of head with dorsal view strongly divergent. long erect and short appressed setae sparsely. Clypeus Petiole in dorsal view with subparallel sides and with two pairs of long setae in anterior portion, one narrow short peduncle anteriorly, distinctly longer than directed upward and the other downward. Anterior wide. Posterior portion of petiole with short process clypeal margin with one pair of long setae medially that is slightly higher than posterior margin of petiole and some pairs of short setae laterally. Scapes with

Figures 53–61. Key to Orthocrema species in Asia. 53–56, propodeal spines: 53, C. macracantha. 54, C. baduvi. 55, C. storki; arrow indicates curved propodeal spine. 56, C. brunensis; arrow indicates straight propodeal spine. 57–58, standing pilosity: 57, C. brevispina. 58, C. quadriruga. 59–61, propodeal spines: 59, C. longipilosa; arrow indicates developed propodeal spine. 60, C. binghamii; arrow indicates undeveloped propodeal spine. 61, C. brevispina; arrow indicates weakly developed propodeal spine. appressed setae. Mesosoma with two pairs of long erect posteriorly. Fourth abdominal tergite with erect setae and stout setae (ps1PN and psaMN) that are much sparsely, but no decumbent to appressed setae. longer than other erect setae. Posterolateral tuber- Body red-brown. Apical two ﬂagellar segments cles of petiole posteriorly with one pair of long yellow, contracting with other ﬂagellar segments that setae. Postpetiole with one pair of long setae on disc are blackish.

Figures 62–70. Key to Orthocrema species in Asia. 62–63, mesosoma sculpture: 62, C. reticulata; 63, C. quadriruga. 64–65, subpetiolar and subpostpetiolar process: 64, C. vieti; left arrow indicates acutely developed subpetiolar process, and right arrow acutely developed subpostpetiolar process. 65, C. quadriruga; left arrow indicates developed subpetiolar process, and right arrow undeveloped subpostpetiolar process. 66–68, compound eyes: 66, C. masukoi; arrow indicates distinctly reduced compound eye. 67, C. ocellata; arrow indicates reduced compound eye. 68, C. biroi; arrow indicates developed compound eye. 69–70, dorsal surface of head: 69, C. reticulata; 70, C. osakensis.

Figures 71–79. Key to Orthocrema species in Asia. 71–72, propodeal spiracles and metapleural gland bulla: 71, C. reticulata; arrows indicate propodeal spiracle apart from metapleural gland bulla. 72, C. bandarensis; arrows indicate propodeal spiracle touching metapleural gland bulla. 73–74, petiole and postpetiole in dorsal view: 73, C. reticulata; arrow indicates petiole without angulate anterolateral corner. 74, C. schimmeri; arrow indicates petiole with angulate anterolateral corner. 75–76, dorsal surface of head: 75, C. biroi; 76, C. fritzi. 77, sculptured surface in front of occipital carinae in C. biroi. 78–79, erect setae on body: 78, C. osakensis; 79, C. fritzi.

Figures 80–87. Key to Orthocrema species in Asia. 80–81, propodeal spines and subpostpetiolar process: 80, C. osakensis; left arrow indicates slender propodeal spine, and right arrow angulate subpostpetiolar process. 81, C. vieti; left arrow indicates thick propodeal spine, and right arrow acute subpostpetiolar process. 82–83, petiole in dorsal view: 82, C. osakensis; dotted lines indicate petiole tapering posteriorly. 83, C. vieti; dotted lines indicate petiole with subparallel sides. 84–85, setae on scape: 84, C. luzonensis; 85, C. fritzi. 86–87, erect setae on anterior and posterior mesonotal ridge: 86, C. luzonensis; arrows indicate erect setae on anterior mesonotal ridge as long as on posterior mesonotal ridge. 87, C. fritzi; arrows indicate erect setae on anterior mesonotal ridge longer than on posterior mesonotal ridge.

Figures 88–95. Key to Orthocrema species in Asia. 88–89, compound eyes: 88, C. myops; arrow indicates distinctly reduced compound eye. 89, C. quadriruga; arrow indicates developed compound eye. 90–91, propodeal spiracles and metapleural gland bulla: 90, C. bandarensis; arrows indicate propodeal spiracle touching metapleural gland bulla. 91, C. quadriruga; arrows indicate propodeal spiracle apart from metapleural gland bulla. 92–93, metanotal groove: 92, C. celebensis; 93, C. philippinensis. 94–95, petiole in dorsal view: 94, C. quadriruga; dotted lines indicate petiole with subparallel sides. 95, C. suehiro; dotted lines indicate petiole tapering anteriorly.

Figures 96–103. Body in lateral view. 96, Crematogaster baduvi, worker [Endau Rompin National Park, W. Malaysia]; 97, Crematogaster bandarensis, worker [Tai Rom Yen National Park, Surat Thani Prov., S. Thailand]; 98, Crematogaster binghamii, worker [Dry evergreen forest, Lumchangwat Stn., Khao Ang Reu Nai WS, Chacheongsap Prov., E. Thai- land]; 99, Crematogaster biroi, worker [Punjavi Univ. Campus, Patiala, Punjab Prov., India]; 100, Crematogaster biroi, intermediate worker [Punjavi Univ. Campus, Patiala, Punjab Prov., India]; 101, Crematogaster brevispina, worker [Tabuc- Tubig, Oriental Dumaguete, Negros, Philippines]; 102, Crematogaster brunensis, worker [Bukit Sulang, nr. Lamunin, Brunei]; 103, Crematogaster celebensis, worker [Kadokaan, P. Batudaka, Togian Island, Sulawesi, Indonesia].

Comments: In the worker this species can be distin- 0.14–0.17; PtW 0.12–0.15; PtH 0.10–0.12; PpL 0.09– guished from all other members of the C. baduvi group 0.11; PpW 0.13–0.16; PtHI 67–82; PtWI 80–88; PpWI by the smooth and shining surfaces of mesosoma, 127–150; WI 100–117. propodeal dorsum (with rugulae), and the very long propodeal spines that are divergent and straight toward General description of worker: Workers monomorphic. tip. This species is similar to C. brunensis, but can be Head subquadratic in full-face view. Mandibles with distinguished from it by the larger head (HW 0.47– four teeth arranged at an equal distance, apical and 0.57, HL 0.48–0.59 vs. 0.43–0.47 and 0.46–0.49 in the subapical teeth large, basal two teeth smaller. Ante- latter), shorter scape (SI 98–108 vs. 113–118) and shorter rior clypeal margin convex in medial portion. Com- propodeal spines (PSL 0.15–0.21 vs. 0.20–0.22). pound eyes slightly projecting beyond lateral margins Specimens from Brunei have the anterior mesonotal of head in full-face view. Scapes reaching posterolateral ridges higher than posterior mesonotal ridges in lateral corners of head. view. Pronotal collar with weakly concave anterior margin in dorsal view, distinctly lower than pronotum in lateral Distribution and biology: This species is known from view. Pronotal dorsum without distinct ridges later- Malaysia (Peninsula), Brunei and Indonesia (Krakatau, ally. Mesonotal dorsum with lateral ridges that ir- Sumatra, Java) (Fig. 49). Specimens from Endau Rompin regularly extend posteriad to tips of propodeal spines. National Park, W. Malaysia and Brunei were collect- Pronotum and mesonotum in lateral view not clearly ed by fogging method. forming continuous dorsal outline; mesonotal dorsum ﬂat. Metanotal groove in dorsal view transverse, almost Material examined: BRUNEI: nine workers, Bukit straight in median portion, forming deep concavity that Sulang, nr. Lamunin, B. M. 1982–388 (fogging), 20.viii is laterally margined by lamellate ridges. Propodeal – 10.ix.1982 (N.E. Stork); INDONESIA: three workers, spiracles elliptical, situated at posterolateral corners Rakata Is. (10 m), Krakatau Isls. (06°09’S 105°28’E), of propodeum, touching to metapleural gland bullae. 11.x.2000, (K. Ogata); one worker, Rakata Is. (20 m), Propodeal spines developed, as long as diameter of Krakatau Isls. (06°09’S 105°28’E), 11.x.2000, (K. Ogata); propodeal spiracles, in dorsal view directed posteriad. three workers, Rakata Is. (20 m), Krakatau Isls. (06°09’S Petiole in dorsal view with subparallel sides, longer 105°28’E), 11.x.2000, (S. Matsui); one worker, Maninjau, than wide. Posterior portion of petiole with short process W. Sumatra, 7–9.viii.1985, (S. & Sk. Yamane) (Muko that is slightly higher than posterior margin of petiole Muko 470 m alt.) (Sumatra Nature Study, SNS Col.); disc in lateral view. Subpetiolar process weakly de- three workers, Jasinga, nr Bogor, W. Java, 5.xi.1985, veloped as angulate tubercle. Postpetiole in lateral view (Sk. Yamane). MALAYSIA: one worker, St. 1, Endau with weakly convex dorsum, as high as petiole, in dorsal Rompin National Park, 6.vii.2003, (H. Kojima et al) view as wide as petiole, weakly bilobed posteriorly but (fogging). without longitudinal sulcus. Subpostpetiolar process undeveloped, but venter of postpetiole convex. Integument essentially smooth and shining. Dorsal CREMATOGASTER BANDARENSIS FOREL, 1913 surface of head smooth and shining. Mandibles with STAT. NOV. feeble rugulae and smooth interspaces. Clypeus gen- (FIG. 97) erally smooth and shining, but with one pair of lon- Crematogaster biroi var. bandarensis Forel, 1913: 76; gitudinal rugulae; rugulae not extending to posterior syntype workers, Bandar Baroe, Sumatra, Indonesia clypeal margin. Dorsal and lateral surfaces of pronotum (v. Buttel-Reepen) (MHNG, examined). One syntype smooth and shining; anterolateral shoulders of pronotum worker (middle specimen of three on one pin) in MHNG with rugulae. Mesopleura smooth and shining. Dorsal here designated Lectotype. surface of propodeum smooth and shining, but one pair Crematogaster biroi var. andelis Santschi 1928: 129. of rugulae running from metanotal groove to tips of One syntype worker, Ile Varela, Pulu Berhala, Sumatra, propodeal spines. Dorsal surface pf petiole smooth. Indonesia, 2.ix.1919 (Corporaal) (NHMB, examined). Lateral surface of petiole generally smooth, but with Syn. nov. one longitudinal rugula. Dorsal and lateral surfaces Crematogaster biroi var. bandarensis; Emery, 1922: of postpetiole smooth and shining. 132 [Combination in C.(Orthocrema)]. Standing pilosity sparse. Dorsal face of head with Crematogaster biroi var. andelis; Stärcke, 1930: 373 suberect setae sparsely. Clypeus with two pairs of long [Description of male]. setae in anterior portion, one directed upward and the other downward. Anterior clypeal margin with one pair Worker measurements (n = 10): HW 0.40–0.47; HL 0.42– of long setae medially and short setae laterally. Scapes 0.50; CI 92–98; SL 0.34–0.39; SI 81–86; EL 0.09– with suberect setae. Mesosoma with ﬁve pairs of long 0.11; PW 0.26–0.29; WL 0.48–0.54; PSL 0.05–0.08; PtL erect and stout setae (ps1PN, ps2PN, psaMN, pspMN,

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 576 S. HOSOISHI AND K. OGATA and ps1PS) that are much longer than other erect setae. Trail, Lambir N. P., Sarawak, E. Malaysia, 2.vii.2004 Posterolateral tubercles of petiole posteriorly with two (SR04-SKY-78) (Sk. Yamane); seven workers, Maliau pairs of long setae. Postpetiole with three pairs of long Basin (riparian forest), Sabah, Borneo (300 m alt.) setae on disc anteriorly, posteriorly and laterally. Fourth 10.xi.2011 (ex carton cover on plant stem with scale abdominal tergite with erect to suberect setae abun- insects) (SB11-SKY-38) (Sk. Yamane); THAILAND: eight dantly, and short decumbent setae sparsely. workers, Khlong Saeng WS, Ratchaprapha D., Surat Body yellow. All ﬂagellar segments yellow. Thani Prov., 15.x.2011 (TH11-SKY-134) (rotting wood) (Sk. Yamane); two workers, Tai Rom Yen NP (400 m Comments: In the worker this species can be distin- alt.), Surat Thani Prov., 11.x.2011 (decayed wood) (TH11- guished from all other members of the C. quadriruga SKY-016) (Sk. Yamane); three workers, Ton Nga Chang, group by the distinct compound eyes, V-shaped nr. Hat Yai, Songkhla Prov., 23.ix.2001 (Eg01-TH- metanotal groove in lateral view, large propodeal spira- 609) (K. Eguchi). cles touching metapleural gland bulla, and short propodeal spines. This species is similar to C. philippinensis and C. sundalandensis, but can be dis- CREMATOGASTER BINGHAMII FOREL, 1904 tinguished from them by the short and stout propodeal (FIG. 98) spines (PSL 0.05–0.08 vs. 0.08–0.11 and 0.08–0.11 in Crematogaster binghamii Forel, 1904: 24; syntype the latters). This species corresponds to sp. 52 of SKY workers, Sikkim, India (MM. Bingham & Moller) (Ito et al., 2001; Eguchi & Yamane, 2003). (MHNG, examined). One syntype worker in MHNG here designated Lectotype. Distribution and biology: This species is known from Crematogaster binghamii; Santschi, 1918: 182 [Com- S. Thailand, Malaysia (Peninsula and Borneo), Brunei, bination in C.(Orthocrema)]. Indonesia (Bali, Sumatra, Krakatau) (Fig. 52). It in- Crematogaster binghamii; Emery, 1922: 131. [Com- habits developed forests, and nests in soil or in leaf bination in C.(Orthocrema)]. litter. Crematogaster binghamii; Blaimer, 2012c: 55 [Combination in C.(Orthocrema)]. Material examined: BRUNEI: one worker, Tasek Merimbun, 16.ii.1999, (Eg99-BOR-126) (K. Eguchi); IN- DONESIA: one worker, Sembung, 25 km N of Denpasar, Worker measurements (n = 10): HW 0.52–0.57; HL 0.52– Bali, 19.iii.1987, (Imai et al); three workers, Tuapejat, 0.59; CI 89–100; SL 0.46–0.50; SI 84–92; EL 0.13– Pulau Sipora, Mentawai Is., W. Sumatra, 24.ii.2007 0.16; PW 0.30–0.34; WL 0.64–0.69; PSL 0–0.03; PtL (SU07-SKY-97) (Sk. Yamane); one worker, Bodong Jaya, 0.18–0.21; PtW 0.16–0.18; PtH 0.12–0.16; PpL 0.13– Sumberjaya, Lampung Barat (secondary forest), S. 0.16; PpW 0.15–0.19; PtHI 67–84; PtWI 80–94; PpWI Sumatra, 16.ix.2007 (leaf litter / s. soil) (Sk. Yamane); 100–138; WI 93–118. six workers, Bodong Java, Sumberjaya, Lampung Barat, S. Sumatra, 16.ix.2007 (leaf litter) (SU07-SKY-156) (Sk. General description of worker: Workers monomorphic. Yamane); four workers, Pulau Sebesi, Lampung Prov., Head subquadratic in full-face view. Mandibles with Sunda Strait, 12.viii.2005 (RK05-SKY-24) (Sk. Yamane); four teeth arranged at an equal distance, apical and MALAYSIA: four workers, 19 miles, Cameron High- subapical teeth large, basal two teeth smaller. Ante- lands, 11.iii.2010 (SH10-Mal-48) (S. Hosoishi); one rior clypeal margin convex in medial portion. Com- worker, Ulu Gombak, Selangor, 27.xi.2005 (MP05-SKY- pound eyes distinctly projecting beyond lateral margins 08) (Sk. Yamane); four workers, UM plantations, Kota of head in full-face view. Scapes reaching posterolateral Tinggi, Johor, 17.xii.2013 (sugar, TD-85) (Sk. Yamane corners of head. & Y. Miyaguni); four workers, UM plantations, Kota Pronotal collar with weakly concave anterior margin Tinggi, Johor, 17.xii.2013 (sugar, GD-64) (Sk. Yamane in dorsal view, distinctly lower than pronotum in lateral & Y. Miyaguni); one worker, Kubah N. P., Kuching, view. Pronotal dorsum without distinct ridges later- Sarawak, Borneo, 28.xii.1997, (F. Yamane); two workers, ally. Mesonotal dorsum with lateral ridges that extend 8 ha plot, Tower Reg., Lambir, Miri, Sarawak #117, posteriad to anterodorsal corners of propodeal dorsum; E Malaysia, 27.viii.1994, (T. Itioka & T. Yumoto); one the ridges forming pair of small triangular processes worker, Old Tower R., Lambir N. P., Miri, Sarawak, (angles) between dorsal and declivity faces of mesonotum 22.i.1993, (Sk. Yamane) (Canopy Ecol.); seven workers, (this condition is more easily seen with mesosoma in Lambir N. P., Miri, Sarawak, 14.v.2011 (SR11-SKY- lateral view). Pronotum and mesonotum in lateral view 11) (nest in soil) (Sk. Yamane); two workers, Danum not clearly forming continuous dorsal outline. Metanotal Valley, Sabah, Borneo, 29.viii.1995, (Sk. Yamane) (prim. groove in dorsal view transverse, almost straight for.); two workers, Tawau Hills N. P., Sabah, Borneo, in median portion, forming deep concavity that is E. Malaysia, 8.vii.1996, (K. Eguchi); three workers, Inoue laterally margined by lamellate ridges. Propodeal

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 577 spiracles oval, situated at posterolateral corners of three workers, NE India, (no date) (S. P. Kurl); NEPAL: propodeum, apart from metapleural gland bullae. three workers, Kathmandu (1350 m alt.), 23.ix.1983 Propodeal spines undeveloped. (M. G. Allen); THAILAND: nine workers, Lumchangwat Petiole in dorsal view with subparallel sides and Stn., Khao Ang Reu Nai WS, Chacheongsao Prov., E. narrow anteriorly, longer than wide. Posterior portion Thailand, 21.viii.2003, (TH03-SKY-55) (Sk. Yamane); of petiole without distinct process in lateral view. VIETNAM: two workers, Rung Thong, Dong Son Dist., Subpetiolar process weakly developed as blunt process. Thanh Hoa Prov. (19°49′S 105°43′E), 30.xi.1999, (K. Postpetiole in lateral view with weakly convex dorsum, Ogata) 15 min TUS #2; one worker, Rung Thong, Dong as high as petiole, in dorsal view as wide as petiole, Son Dist., Thanh Hoa Prov. (19°49′S 105°43′E), globular, not bilobed. Subpostpetiolar process unde- 1.xii.1999, (K. Ogata) 15 min TUS #2. veloped, but venter of postpetiole convex. Integument essentially smooth and shining. Dorsal surface of head smooth and shining. Mandibles with CREMATOGASTER BIROI MAYR, 1897 feeble rugulae and smooth interspaces. Clypeus gen- (FIGS 99, 100) erally smooth and shining, but with one to two dis- Crematogaster biroi Mayr, 1897: 428; syntype workers, tinct pairs of longitudinal rugulae; rugulae not extending Columbo, Sri Lanka (Biró) (HNHM, examined). One to posterior clypeal margin. Dorsal and lateral sur- syntype worker (with label of 160) in HNHM from faces of pronotum smooth and shining; anterolateral Columbo here designated Lectotype. shoulders of pronotum with rugulae. Mesopleura gen- Crematogaster biroi var. aitkenii Forel, 1902a: 203; erally smooth and shining. Rugula on higher portion syntype workers, Kanara, India (Aitken) (MHNG, ex- of mesopleura extending to small pit of mesothoracic amined). Syn. nov. One syntype worker (top speci- spiracles. Dorsal surface of propodeum smooth and men of three on one pin) in MHNG here designated shining. Dorsal surface of petiole smooth and shining. Lectotype. Lateral surface of petiole generally smooth, but with Crematogaster biroi var. smythiesii Forel, 1902a: 203; one longitudinal rugula. Dorsal and lateral surfaces syntype workers, Dehra Dun, India (Smythies) (MHNG, of postpetiole smooth and shining. examined). Syn. nov. One syntype worker (top speci- Standing pilosity abundant. Dorsal face of head with men of two on one pin) in MHNG here designated erect to suberect setae abundantly. Clypeus with two Lectotype. pairs of long setae in anterior portion, one directed Crematogaster urvijae Bharti, 2003: 85, ﬁgs 1–9; upward and the other downward. Anterior clypeal holotype and paratype workers, Patiala, Punjab, India, margin with one pair of long setae medially and short 250 m alt., 7.June.1999 (H. Bharti) (examined). Syn. setae laterally. Scapes with suberect setae. Mesosoma nov. with seven pairs of long erect and stout setae (ps1PN, Crematogaster biroi; Bingham, 1903: 138 [Descrip- ps2PN, psaMN, pspMN, psPR, ps1PS, and ps2PS) that tion of intermediate worker]. are much longer than other erec setae. Posterolateral Crematogaster biroi; Emery, 1922: 131 [Combina- tubercles of petiole posteriorly with three pairs of long tion in C.(Orthocrema)]. setae. Postpetiole with three pairs of long setae on disc Crematogaster biroi var. aitkenii; Emery, 1922: 132 anteriorly, posteriorly and laterally. Fourth abdomi- [Combination in C.(Orthocrema)]. nal tergite with erect to suberect setae abundantly, but Crematogaster biroi var. smythiesii; Emery, 1922: 132 no decumbent to appressed setae. [Combination in C.(Orthocrema)]. Body yellow. All ﬂagellar segments yellow. Crematogaster biroi; Imai et al. 1984: 6 [Karyotype]. Comments: In the worker this species can be distin- Crematogaster aitkenii; Wu & Wang, 1992: 1319 guished from all other members of the C. binghamii [Raised to species]. group by the undeveloped or weakly developed propodeal Crematogaster biroi; Blaimer, 2012c: 55 [Combina- spines. This species is similar to C. brevispina, but can tion in C.(Orthocrema)]. be distinguished from it by the weakly developed Crematogaster urvijae; Blaimer, 2012c: 55 [Combi- propodeal spines (PSL 0–0.03 vs. 0.04–0.07 in the nation in C.(Crematogaster)]. latter); the length smaller than the diameter of propodeal spiracles. Worker measurements (n = 10): HW 0.44–0.55; HL 0.45– Distribution: This species is known from Nepal, India, 0.52; CI 98–106; SL 0.33–0.39; SI 70–75; EL 0.09– Bangladesh, Thailand and Vietnam (Fig. 50). 0.13; PW 0.30–0.34; WL 0.49–0.58; PSL 0.09–0.12; PtL 0.16–0.19; PtW 0.13–0.17; PtH 0.12–0.15; PpL 0.10– Material examined: BANGLADESH: three workers, 0.13; PpW 0.14–0.17; PtHI 72–82; PtWI 78–89; PpWI Jointapur (30 min sample), 30.x.1994, (K. Ogata); INDIA: 125–141; WI 100–108.

General description of worker: Workers monomorphic, Standing pilosity sparse. Dorsal face of head with but intermediate worker as large as queen (details several pairs (six to seven) of erect and stout setae, below). and short and appressed setae abundantly. Clypeus with Head subquadratic in full-face view. Mandibles with two pairs of long setae in anterior portion, one direct- four teeth arranged at an equal distance, apical and ed upward and the other downward. Anterior clypeal subapical teeth large, basal two teeth smaller. Ante- margin with two pairs of long setae medially and some rior clypeal margin convex in medial portion. Com- pairs (three to four) of short setae laterally. Scapes with pound eyes distinctly projecting beyond lateral margins suberect setae. Mesosoma with four pairs of long erect of head in full-face view. Scapes reaching posterolateral and stout setae (ps1PN, psaMN, pspMN, and ps1PS) corners of head. that are much longer than other erect setae. Pronotal collar with almost straight anterior margin Posterolateral tubercles of petiole posteriorly with one in dorsal view, distinctly lower than pronotum in lateral pair of stout setae. Postpetiole with one pair of stout view. Pronotal dorsum with feeble ridges laterally. setae on disc posteriorly. Fourth abdominal tergite with Mesonotal dorsum with lateral ridges that irregular- three or six pairs of erect and stout setae, and short ly extend posteriad to tips of propodeal spines. Pronotum appressed setae abundantly. and mesonotum in lateral view forming slightly convex, Body yellow. All ﬂagellar segments yellow. continuous dorsal outline. Metanotal groove in dorsal view transverse, almost straight in median portion, Intermediate worker measurements (n = 1): HW 0.82; forming deep concavity that is laterally margined by HL 0.79; CI 104; SL 0.48; SI 59; EL 0.21; PW 0.58; lamellate ridges. Propodeal spiracles oval, situated WL 0.98; PSL 0.18; PtL 0.35; PtW 0.28; PtH 0.25; at posterolateral corners of propodeum, apart PpL 0.21; PpW 0.32; PtHI 71; PtWI 80; PpWI 152; WI from metapleural gland bullae. Propodeal spines 114. developed, longer than diameter of propodeal spiracles, in dorsal view directed posteriad. Description of intermediate worker: With worker char- Petiole in dorsal view with parallel sides and angulate acter conditions, except as follows. shoulders anteriorly, longer than wide. Posterior portion Head subquadratic. of petiole with short process that is slightly higher than Mesonotum highly convex in lateral view. Mesonotal posterior margin of petiole disc in lateral view. dorsum without lateral ridges. Pronotum forming same Subpetiolar process strongly developed as acute process. dorsal outline with mesonotum in lateral view, but pos- Postpetiole in lateral view with weakly convex dorsum, terior face forming vertical slope to metanotal groove. as high as petiole, in dorsal view as wide as or slight- Propodeal spiracles elliptical. ly wider than petiole, weakly bilobed posteriorly but Subpostpetiolar area not observable in the speci- without longitudinal sulcus. Subpostpetiolar process men examined. developed as process. Clypeus with some longitudinal rugulae: rugulae ex- Integument essentially sculptured. Dorsal surface of tending to frontal region between frontal carinae. head generally smooth on central region, but weakly Dorsal face of head with erect setae sparsely. Clypeus sculptured reticulately laterally. Occipital region near with one pair of long setae around antennal bases; di- margin weakly sculptured. Mandibles with feeble rected upper. Mesosoma without distinct erect setae. rugulae and smooth interspaces. Clypeus generally Fourth abdominal tergite with short decumbent to smooth, but with two distinct pairs of longitudinal appressed setae sparsely. rugulae and weakly sculptured interspaces; rugulae not extending to posterior clypeal margin. Anterolateral Comments: In the worker this species is very distinct shoulders of pronotum with rugulae. Pronotum and among the C. biroi group in having the sculptured mesonotum with longitudinal rugulae and sculp- surface near occipital carinae of head, the strongly de- tured interspaces. Lateral surface of pronotum smooth veloped subpetiolar process, and acutely developed and shining on central portion, but weakly sculp- subpostpetiolar process. tured on surrounding. Mesopleura weakly sculptured, but relatively smooth in central areas. Rugula Distribution and biology: This species is known from on higher portion of mesopleura extending to small pit India and Sri Lanka (Fig. 51). Although this species of mesothoracic spiracles. One pair of rugulae running has been reported from China by Wu & Wang (1995) from metanotal groove to tips of propodeal spines and Zhou (2001), and from Taiwan by Terayama (2009), (rugulae on mesonotum extending posteriorly to we have not been able to examine those specimens. propodeal spines). Dorsal surface of propodeum weakly This species nests in soil. sculptured. Dorsal and lateral surfaces of petiole sculptured. Dorsal surface of postpetiole smooth and shining. Material examined: INDIA: two workers, Chandi- Lateral surface of postpetiole weakly sculptured. garh, Punjab, 10.viii.1978, (H. Imai et al); ten workers,

Punjavi Univ. Campus, Patiala, Punjab Prov., 14.x.2007 tinct pairs of longitudinal rugulae; longer rugulae not (open site) (IN07-SKY-019) (Sk. Yamane). extending to posterior clypeal margin. Costulate rugulae weakly developed on malar region. Dorsal and lateral surfaces of pronotum smooth and shining; anterolateral CREMATOGASTER BREVISPINA SP. NOV. shoulders of pronotum without rugulae. Mesopleura (FIG. 101) generally smooth and shining. Rugula on higher portion Holotype worker. Tabuc-Tubig, Oriental Dumaguete, of mesopleura extending to small pit of mesothoracic Negros, PHILIPPINES, 27.v.1983 (707) (C. K. Stall & spiracles. Dorsal surface of propodeum generally smooth F. P. Godoy) (BMNH). and shining. Dorsal surface of petiole smooth and Paratypes. Eight workers, same data as holotype shining. Lateral surface of petiole generally smooth, (BMNH, CASC, KUEC, MHNG, MPMP, THNHM). but with one longitudinal rugula running from spiracles to posterior margin. Dorsal and lateral surfaces Worker measurements (n = 9): HW 0.51–0.62; HL 0.54– of postpetiole smooth and shining. 0.65; CI 93–98; SL 0.50–0.55; SI 92–96; EL 0.15– Standing pilosity abundant. Dorsal face of head with 0.17; PW 0.34–0.41; WL 0.61–0.72; PSL 0.04–0.07; PtL erect to suberect setae abundantly. Clypeus with three 0.19–0.25; PtW 0.17–0.21; PtH 0.13–0.17; PpL 0.13– pairs of long setae on anterior portion, one directed 0.15; PpW 0.17–0.21; PtHI 65–70; PtWI 74–90; PpWI upward, one downward, the other laterally below 120–140; WI 95–106. antennal sockets. Anterior clypeal margin with single long setae medially and one pair of long setae later- General description of worker: Workers monomorphic. ally, and short setae laterally. Gena (malar space) with Head subquadratic in full-face view. Mandibles some suberect setae near mandibular insertion. Scapes with four teeth arranged at an equal distance, apical with suberect to decumbent setae; short setae basally and subapical teeth large, basal two teeth smaller. and long setae distally. Mesosoma with seven to eight Anterior clypeal margin convex in medial portion. Com- distinct pairs of long erect and stout setae (ps1PN, pound eyes distinctly projecting beyond lateral margins ps2PN, psaMN, pspMN, psPR, ps1PS, and one to two of head in full-face view. Scapes reaching posterolateral ps2PS) that are much longer than other erect setae. corners of head. Posterolateral tubercles of petiole posteriorly with Pronotal collar with concave anterior margin in dorsal three pairs of long setae. Postpetiole with four pairs view, distinctly lower than pronotum in lateral view. of long setae on disc anterodorsally, anterolaterally, Pronotal dorsum without distinct ridges laterally. posteromedially and posteriorly. Fourth abdominal Mesonotal dorsum with lateral ridges; the ridges forming tergite with erect to suberect setae abundantly, but no pair of small triangular processes (angles) between decumbent to appressed setae. dorsal and declivity faces of mesonotum (this condi- Body yellow. All ﬂagellar segments yellow. tion is more easily seen with mesosoma in lateral view). Pronotum and mesonotum in lateral view not clearly Comments: In the worker this species can be distin- forming continuous dorsal outline. Metanotal groove guished from all other members of the C. binghamii in dorsal view transverse, almost straight in median group by the weakly developed propodeal spines. This portion, forming deep concavity that is laterally mar- species is similar to C. binghamii, but can be distin- gined by lamellate ridges. Propodeal spiracles oval, situ- guished from it by the propodeal spines (PSL 0.04– ated at posterolateral corners of propodeum, apart from 0.07 vs. 0–0.03 in the latter); the length as large as metapleural gland bullae. Propodeal spines weakly de- the diameter of propodeal spiracles. veloped, shorter than diameter of propodeal spiracles, in dorsal view directed posteriad. Distribution: This species is known from the type lo- Petiole in dorsal view with subparallel sides and cality of the Philippines (Fig. 50). narrow anteriorly, longer than wide. Posterior portion of petiole without distinct process in lateral view. Etymology: The speciﬁc name refers to the short Subpetiolar process weakly developed as blunt process. propodeal spines. Postpetiole in lateral view with weakly convex dorsum, as high as petiole, in dorsal view as wide as petiole, CREMATOGASTER BRUNENSIS SP. NOV. globular, not bilobed. Subpostpetiolar process undeveloped, but venter of postpetiole weakly convex. (FIG. 102) Integument essentially smooth and shining. Dorsal Holotype worker. nr. Lamunin, Bukit Sulang, BRUNEI, surface of head smooth and shining, but with rugulae B. M. 1982-388 Fogging, 20.viii–10.ix.1982 (N. E. Stork) on surrounding region of antennal sockets. Mandi- (BMNH). bles with feeble rugulae and smooth interspaces. Clypeus Paratypes. Five workers, same data as holotype generally smooth and shining, but with one to two dis- (CASC, ITBC, KUEC, MHNG, THNHM).

Worker measurements (n = 6): HW 0.43–0.47; HL 0.46– ward. Anterior clypeal margin with one pair of long 0.49; CI 92–100; SL 0.51–0.53; SI 113–118; EL 0.12– setae medially and some pairs of short setae lateral- 0.13; PW 0.26–0.29; WL 0.60–0.63; PSL 0.20–0.22; PtL ly. Scapes with suberect to decumbent setae. Mesosoma 0.23–0.26; PtW 0.14–0.17; PtH 0.14–0.15; PpL 0.12– with three pairs of long erect and stout setae (ps1PN, 0.13; PpW 0.15–0.17; PtHI 56–61; PtWI 60–67; PpWI psaMN, and pspMN) that are much longer than other 123–133; WI 100–107. erect setae. Posterolateral tubercles of petiole posteriorly with one pair of long setae. Postpetiole with two General description of worker: Workers monomorphic. pairs of long setae on disc anteriorly, posteriorly. Fourth Head round in full-face view. Mandibles with four abdominal tergite with erect setae sparsely, but no de- teeth arranged at an equal distance, apical and cumbent to appressed setae. subapical teeth large, basal two teeth smaller. Ante- Body red-brown. Apical two flagellar segments light rior clypeal margin convex in medial portion. Com- yellow, contracting with other flagellar segments that pound eyes distinctly projecting beyond lateral margins are yellow. of head in full-face view. Scapes exceeding posterolateral corners of head. Comments: In the worker this species can be distin- Pronotal collar with concave anterior margin in dorsal guished from all other members of the C. baduvi group view, slightly lower than pronotum in lateral view. by the smooth and shining surface of mesosoma, smooth Pronotal dorsum without distinct ridges laterally. surface of propodeal dorsum, and propodeal spines de- Mesonotal dorsum with lateral ridges that irregular- veloped straight at the tip. This species is similar to ly extend posteriad to tips of propoedal spines. Pronotum C. baduvi, but can be distinguished from it by the and mesonotum in lateral view forming evenly arched, smaller head (HW 0.43–0.47, HL 0.46–0.49 vs. 0.47– continuous dorsal outline. Metanotal groove in dorsal 0.57 and 0.48–0.59 in the latter) and longer propodeal view transverse, almost straight in median portion, spines (PSL 0.20–0.22 vs. 0.15–0.21). forming shallow concavity that is laterally margined by ridges. Propodeal spiracles oval, situated at Distribution and biology: This species is known only posterolateral corners of propodeum, apart from from the type locality in Brunei (Fig. 49). Type speci- metapleural gland bullae. Propodeal spines devel- mens were collected by fogging method. oped, longer than diameter of propodeal spiracles, in dorsal view strongly divergent. Etymology: The specific name refers to the country of Petiole in dorsal view with subparallel sides and origin, Brunei. narrow short peduncle anteriorly, distinctly longer than wide. Posterior portion of petiole with short process that is slightly higher than posterior margin of petiole CREMATOGASTER CELEBENSIS SP. NOV. disc in lateral view. Subpetiolar process undeveloped. (FIG. 103) Postpetiole in lateral view with strongly convex dorsum, distinctly higher than petiole, in dorsal view as wide Holotype worker. Kadokaan, P. Batudaka, Togian Island, as petiole, globular, not bilobed. Subpostpetiolar process Sulawesi, INDONESIA, 23.viii.2008 (CE08-SKY-14) (Sk. undeveloped, but venter of postpetiole convex. Yamane) (MBBJ). Integument essentially smooth and shining. Dorsal Paratypes. Three workers, same data as holotype surface of head smooth and shining. Mandibles with (BMNH, CASC, KUEC). feeble rugulae and smooth interspaces. Clypeus generally smooth and shining, but with one pair of lon- Worker measurements (n = 5): HW 0.49–0.51; HL 0.51– gitudinal rugulae; rugulae not extending to the posterior 0.53; CI 94–96; SL 0.39–0.42; SI 80–84; EL 0.10– clypeal margin. Dorsal and lateral surfaces of pronotum 0.11; PW 0.31–0.33; WL 0.56–0.59; PSL 0.09–0.10; PtL smooth and shining; anterolateral shoulders of pronotum 0.19–0.20; PtW 0.15–0.17; PtH 0.13–0.15; PpL 0.11– with feeble rugulae. Mesopleura generally sculptured, 0.13; PpW 0.16–0.18; PtHI 68–75; PtWI 79–89; PpWI but sometimes smooth except for their marginal areas 133–145; WI 100–107. in some specimens. Dorsal surface of propodeum generally smooth and shining, but with rugulae dorsolateral General description of worker: Workers monomorphic. areas. Dorsal surface of petiole smooth and shining. Head subquadratic in full-face view. Mandibles with Lateral surface of petiole weakly sculptured. Dorsal four teeth arranged at an equal distance, apical and and lateral surfaces of postpetiole smooth and shining. subapical teeth large, basal two teeth smaller. Ante- Standing pilosity sparse. Dorsal face of head with rior clypeal margin convex in medial portion. Com- long erect (three pairs) and short appressed setae sparse- pound eyes slightly projecting beyond lateral margins ly. Clypeus with two pairs of long setae in anterior of head in full-face view. Scapes reaching posterolateral portion, one directed upward and the other down- corners of head.

Pronotal collar with weakly concave anterior margin Comments: In the worker this species can be distin- in dorsal view, distinctly lower than pronotum in lateral guished from all other members of the C. quadriruga view. Pronotal dorsum without distinct ridges later- group by the distinct compound eyes, U-shaped ally. Mesonotal dorsum with lateral ridges that ir- metanotal groove in lateral view, large propodeal spira- regularly extend posteriad to tips of propodeal spines. cles touching metapleural gland bulla, and longer Pronotum and mesonotum in lateral view forming slight- propodeal spines. This species is similar to C. ly convex, continuous dorsal outline. Metanotal groove banadarensis, but can be distinguished from it by the in dorsal view transverse, almost straight in median deep (or U-shaped) metanotal groove in lateral view portion, forming deep concavity that is laterally mar- and long propodeal spines (PSL 0.09–0.10 vs. 0.05– gined by lamellate ridges (U-shaped in lateral view). 0.08 in the latter). Propodeal spiracles elliptical, situated at posterolateral corners of propodeum, touching to metapleural gland Distribution: This species is known from Indonesia bullae. Propodeal spines developed, as long as diam- (Sulawesi) (Fig. 52). eter of propodeal spiracles, in dorsal view directed posteriad. Etymology: The specific name refers to the former name Petiole in dorsal view with subparallel sides and of Sulawesi, Celebes. narrow anteriorly, longer than wide. Posterior portion of petiole with short process that is slightly higher than Material examined: INDONESIA: one worker, Tanimpo, posterior margin of petiole disc in lateral view. P. Batudaka, Togian Is., S. Sulawesi, 23.viii.2008 (Sk. Subpetiolar process undeveloped. Postpetiole in lateral Yamane); nine workers, Ranu River Area, nr. Morowali, view with weakly convex dorsum, as high as petiole, Tengah, Sulawesi, 27.i.-20.iv.1980 (B.M. 1980-280) in dorsal view as wide as petiole, weakly bilobed (M.J.D. Brendell). posteriorly but without longitudinal sulcus. Subpostpetiolar process undeveloped, but venter of postpetiole convex. CREMATOGASTER FRITZI EMERY, 1901 Integument essentially smooth and shining. Dorsal (FIG. 104) surface of head smooth and shining. Clypeus generally smooth and shining, but with one distinct pair of Crematogaster fritzi Emery, 1901: 576, fig. B; syntype longitudinal rugulae; rugulae not extending to pos- workers, Tomohon, Sulawesi, Indonesia (MCSN, ex- terior clypeal margin. Dorsal and lateral surfaces of amined). One syntype worker (basal specimen of three pronotum smooth and shining; anterolateral shoul- of top label of one pin) in MCSN here designated ders of pronotum with rugulae. Mesopleura smooth and Lectotype. shining. Dorsal surface of propodeum generally smooth Crematogaster sordidula r. fritzi; Forel, 1902b: 410 and shining, but one pair of rugulae running from [Race of sordidula]. metanotal groove to tips of propodeal spines. Dorsal Crematogaster fritzi; Emery, 1912: 668 [Revived status surface of petiole smooth and shining. Lateral surface as species]. of petiole generally smooth, but with one longitudi- Crematogaster fritzi; Emery, 1922: 132 [Combina- nal rugula. Dorsal and lateral surfaces of postpetiole tion in C.(Orthocrema)]. smooth and shining. Crematogaster fritzi; Blaimer, 2012c: 55 [Combina- Standing pilosity sparse. Dorsal face of head with tion in C.(Orthocrema)]. suberect setae sparsely. Mandibles with feeble rugulae and smooth interspaces. Clypeus with two pairs of Worker measurements (n = 14): HW 0.40–0.48; HL 0.42– long setae in anterior portion, one directed upward 0.49; CI 95–100; SL 0.34–0.40; SI 80–89; EL 0.10– and the other downward. Anterior clypeal margin 0.13; PW 0.24–0.31; WL 0.45–0.54; PSL 0.07–0.1; PtL with one pair of long setae medially and short 0.15–0.18; PtW 0.13–0.16; PtH 0.11–0.13; PpL 0.09– setae laterally. Scapes with suberect setae. Mesosoma 0.12; PpW 0.14–0.17; PtHI 69–76; PtWI 82–94; PpWI with five pairs of long erect and stout setae (ps1PN, 127–167; WI 100–108. ps2PN, psaMN, pspMN, and ps1PS) that are much longer than other erect setae. Posterolateral tuber- General description of worker: Workers monomorphic. cles of petiole posteriorly with three pairs of stout Head subquadratic in full-face view. Mandibles with setae (one pair of shorter setae laterally). Postpetiole four teeth arranged at an equal distance, apical and with three pairs of long setae on disc anteriorly, pos- subapical teeth large, basal two teeth smaller. Ante- teriorly and laterally. Fourth abdominal tergite with rior clypeal margin convex in medial portion. Com- erect to suberect setae abundantly, and short decum- pound eyes distinctly projecting beyond lateral margins bent setae sparsely. of head in full-face view. Scapes reaching posterolateral Body yellow. All flagellar segments yellow. corners of head.

Figures 104–111. Body in lateral view. 104, Crematogaster fritzi, worker [Bukit Larut (Maxwell’s Hill), Perak, W. Ma- laysia]; 105, Crematogaster longipilosa, worker [7 miles, Cameron Highlands, W. Malaysia]; 106, Crematogaste luzonensis, worker [Luzon, Philippines]; 107, Crematogaster macracantha, worker [Poring, Sabah, Borneo, E. Malaysia]; 108, Crematogaster masukoi, worker [Sepilok, Sandakan, Borneo, E. Malaysia]; 109, Crematogaster moatensis, worker [nr. Kotamobaqu, Danau Mooat, Utara, Sulawesi, Indonesia]; 110, Crematogaster myops, worker [Old Tower Region, Lambir National Park, Sarawak, Borneo, E. Malaysia]; 111, Crematogaster ocellata, worker [Tu Lung, Mai Chau Dist., Hoa Binh Prov., N. Vietnam].

Pronotal collar with almost straight anterior margin abdominal tergite with three to four pairs of erect and in dorsal view, distinctly lower than pronotum in lateral stout setae, and short appressed setae abundantly. view. Pronotal dorsum with feeble ridges laterally. Body yellow. All flagellar segments yellow. Mesonotal dorsum with lateral ridges that irregular- Comments: In the worker this species can be distin- ly extend posteriad to tips of propodeal spines. Pronotum guished from all other members of the C. biroi group and mesonotum in lateral view forming slightly convex, by the scape with appressed setae, distinct com- continuous dorsal outline. Metanotal groove in dorsal pound eyes, generally smooth dorsal surface of head, view transverse, almost straight in median portion, and sparse erect and stout setae on body. This species forming deep concavity that is laterally margined by is similar to C. luzonensis, but can be distinguished lamellate ridges. Propodeal spiracles oval, situated at from it by the appressed setae on scape, longer setae posterolateral corners of propodeum, apart from on anterior mesonotal ridges, sparse erect setae on metapleural gland bullae. Propodeal spines devel- fourth abdominal tergite. oped, longer than diameter of propodeal spiracles, in dorsal view directed posteriad. Distribution and biology: This species is known from Petiole in dorsal view with subparallel sides and Malaysia (Peninsula), Brunei, Indonesia (Sumatra, narrow anteriorly, longer than wide. Posterior portion Sulawesi) (Fig. 51). This species inhabits developed of petiole with short process that is slightly higher than forests, and nests in dead twigs on trees. posterior margin of petiole disc in lateral view, with Material examined: BRUNEI: five workers, nr Lamunin, node-like process in lateral view. Subpetiolar process Bukit Sulang, 20. viii. – 10. ix. 1982 (B. M. 1982- developed as blunt process. Postpetiole in lateral view 388. Fogging) (N. E. Stork); INDONESIA: three workers, with weakly convex dorsum, as high as petiole, in dorsal Babulmakmur, near Sibigo, Pulau Simeulue, Aceh, view as wide as or slightly narrower than petiole, weakly Sumatra, 15. ix. 2012 (decayed stem) (SU12-SKY- bilobed posteriorly but without longitudinal sulcus. 091) (Sk. Yamane); two workers, Lamerem, Pulau Subpostpetiolar process developed bluntly. Simeulue, Aceh, Sumatra, 13. ix. 2012 (nest on tree Integument essentially sculptured. Dorsal surface of trunk) (SU12-SKY-068) (Sk. Yamane); three workers, head generally smooth, but with rugulae on surround- Dumoga-Bone N. P., Utara, Sulawesi (400 m) (Fog. 5) ing region of antennal sockets. Mandibles with feeble (BMNH Plot C), 11. ii. 1985 (N. Stork); MALAYSIA: rugulae and smooth interspaces. Clypeus generally 14 workers, Bukit Larut (Maxwell’s Hill), 4. x. 2011, smooth, but with one distinct pair of longitudinal (SH11-Mal-13) (dead twig on tree) (S. Hosoishi); seven rugulae and one pair of feeble rugulae and weakly sculp- workers, Setiu Wetland, Besut, Terengganu, 13. xii. 2010 tured interspaces; distinct rugulae extending to pos- (dead twig on tree) (MP10-SKY-08) (Sk. Yamane). terior clypeal margin. Anterolateral shoulders of pronotum with rugulae. Dorsal surface of pronotum CREMATOGASTER GAVAPIGA MENOZZI, 1935 with longitudinal rugulae and weakly sculptured Crematogaster (Orthocrema) gavapiga Menozzi, 1935: interspaces. Lateral surface of pronotum generally 109, fig. 5; worker, Tjampea, Java, Indonesia (2389) smooth and shining. Mesopleura weakly sculptured. (probably in IEGG, not seen). Pronotum and mesonotum with longitudinal rugulae Crematogaster gavapiga; Blaimer, 2012c: 55 [Com- and sculptured interspaces. Dorsal surface of propodeum bination in C.(Orthocrema)]. generally smooth and shining, but one pair of rugulae running from metanotal groove extending posteriorly Comments: We have not been able to examine the types and diverging to tips of propodeal spines. Dorsal surface of C. gavapiga. From the original description and key of petiole generally smooth. Lateral surface of petiole provided by Menozzi (1935), this species is close to C. sculptured. Dorsal and lateral surfaces of postpetiole bandarensis. He distinguished C. gavapiga from C. smooth and shining. bandarensis by smooth and shining surface of dorsal Standing pilosity sparse. Dorsal face of head with mesosoma. However, some specimens in our collec- three pairs of erect and stout setae, and short and tions of C. bandarensis have smooth and shining surface appressed seate abundantly. Clypeus with two pairs of dorsal mesosoma, suggesting that the character state of long and stout setae in anterior portion, one direct- seen in C. gavapiga is within the infraspecific varied upward and the other downward. Anterior clypeal ation of C. bandarensis. Taxonomic relationship with margin with one pair of long setae medially and some C. bandarensis will remain uncertain until type ma- pair of short setae laterally. Scapes with appressed setae. terial can be examined. Mesosoma with three pairs of long erect and stout setae (ps1PN, psaMN, and pspMN) that are much longer than CREMATOGASTER JAVANICA MENOZZI, 1935 other erect setae. Posterolateral tubercles of petiole pos- Crematogaster (Orthocrema) javanica Menozzi, 1935: teriorly with one pair of stout setae. Postpetiole with 108, fig. 4; worker, Tjiapers, Java, Indonesia (2365) two pairs of stout setae on disc posteriorly. Fourth (probably in IEGG, not seen).

Crematogaster javanica; Hosoishi et al. 2010: 346 diameter of propodeal spiracles, in dorsal view direct- [Taxonomic remark]. ed posteriad. Crematogaster javanica; Blaimer, 2012c: 55 [Com- Petiole in dorsal view with subparallel sides and bination in C.(Orthocrema)]. narrow anteriorly, longer than wide. Posterior portion of petiole without distinct process in lateral view. Comments: This species may be a close relative of C. Subpetiolar process weakly developed as blunt process. myops (Hosoishi et al., 2010). Postpetiole in lateral view with weakly convex dorsum, as high as petiole, in dorsal view as wide as petiole, globular, not bilobed. Subpostpetiolar process unde- CREMATOGASTER LONGIPILOSA FOREL, 1907 veloped, but venter of postpetiole weakly convex. (FIG. 105) Integument essentially smooth and shining. Dorsal surface of head smooth and shining, but with rugulae Crematogaster longipilosa Forel, 1907: 24; syntype on surrounding region of antennal sockets. Mandi- workers, Kuala Lumpur, Malaysia (Biró) (MHNG, ex- bles with feeble rugulae and smooth interspaces. Clypeus amined). One syntype worker (top specimen of two on generally smooth and shining, but with one to two dis- one pin) in MHNG here designated Lectotype. tinct pairs of longitudinal rugulae; rugulae not ex- Crematogaster longipilosa; Forel, 1911b: 383 [De- tending to posterior clypeal margin. Costulate rugulae scriptions of queen and male]. present on malar region. Dorsal and lateral surfaces Crematogaster longipilosa; Viehmeyer, 1916: 124 of pronotum smooth and shining; anterolateral shoul- [Description of queen]. ders of pronotum without rugulae. Mesopleura gen- Crematogaster longipilosa; Emery, 1922: 132 [Com- erally smooth and shining. Rugula on higher portion bination in C.(Orthocrema)]. of mesopleura extending to small pit of mesothoracic Crematogaster longipilosa; Blaimer, 2012c: 55 [Com- spiracles. Dorsal surface of propodeum generally smooth bination in C.(Orthocrema)]. and shining, but few longitudinal rugulae on anteri- or areas. Dorsal surface of petiole smooth and shining. Worker measurements (n = 10): HW 0.60–0.69; HL 0.63– Lateral surface of petiole generally smooth, but with 0.71; CI 95–97; SL 0.52–0.58; SI 84–89; EL 0.15– one longitudinal rugula running from spiracles to pos- 0.18; PW 0.37–0.40; WL 0.71–0.80; PSL 0.13–0.16; PtL terior margin. Dorsal and lateral surfaces of postpetiole 0.25–0.29; PtW 0.20–0.23; PtH 0.16–0.19; PpL 0.16– smooth and shining. 0.19; PpW 0.20–0.24; PtHI 62–73; PtWI 79–92; PpWI Standing pilosity abundant. Dorsal face of head with 111–137; WI 95–105. erect to suberect setae abundantly. Clypeus with three pairs of long setae on anterior portion, one directed General description of worker: Workers monomorphic. upward, one downward. the other laterally below Head subquadratic in full-face view. Mandibles with antennal sockets. Anterior clypeal margin with single four teeth arranged at an equal distance, apical and long setae medially and one pair of long setae later- subapical teeth large, basal two teeth smaller. Ante- ally, and short setae laterally. Gena (malar space) with rior clypeal margin convex in medial portion. Com- some suberect setae near mandibular insertion. Scapes pound eyes distinctly projecting beyond lateral margins with suberect to decumbent setae; short setae basally of head in full-face view. Scapes reaching posterolateral and long setae distally. Mesosoma with seven to eight corners of head. distinct pairs of long erect and stout setae (ps1PN, Pronotal collar with weakly concave anterior margin ps2PN, psaMN, pspMN, psPR, ps1PS, and one to two in dorsal view, distinctly lower than pronotum in lateral ps2PS) that are much longer than other erect setae. view. Pronotal dorsum without distinct ridges later- Posterolateral tubercles of petiole posteriorly with three ally. Mesonotal dorsum with lateral ridges that ir- pairs of long setae. Postpetiole with four pairs of long regularly extend posteriad to tips of propodeal spines; setae on disc anterodorsally, anterolaterally, the ridges forming pair of small triangular processes posteromedially and posteriorly. Fourth abdominal (angles) between dorsal and declivity faces of mesonotum tergite with erect to suberect setae abundantly, but no (this condition is more easily seen with mesosoma in decumbent to appressed setae. lateral view). Pronotum and mesonotum in lateral view Body yellow to brown. All ﬂagellar segments yellow. not clearly forming continuous dorsal outline. Metanotal groove in dorsal view transverse, almost straight in Comments: In the worker this species is very distinct median portion, forming deep concavity that is later- among the C. binghamii group in having developed ally margined by lamellate ridges. Propodeal spira- propodeal spines (PSL 0.13–0.16 vs. 0–0.07 in the cles oval, situated at posterolateral corners of others). propodeum, apart from (or slightly touching) metapleural This species corresponds to C. sp. 51 of SKY (Ito et al., gland bullae. Propodeal spines developed, longer than 2001).

Distribution and biology: This species is known from pound eyes distinctly projecting beyond lateral margins S. Thailand, Malaysia (Peninsula) and Indonesia of head in full-face view. Scapes reaching posterolateral (Kalimantan, Java, Sumatra) (Fig. 50). This species in- corners of head. habits disturbed to developed forests, and nests in soil. Pronotal collar with almost straight anterior margin in dorsal view, distinctly lower than pronotum in lateral Material examined: INDONESIA: two workers, view. Pronotal dorsum with feeble ridges laterally. Sangkimah, Kutai N. P., E. Kalimantan, 13.ix.1993 Mesonotal dorsum with lateral ridges that irregular- (roadside) (Sk. Yamane); one worker, Kutai, Kalimantan, ly extend posteriad to tips of propodeal spines. Pronotum 17.viii.1986 (T. Yajima); six workers, Gn Salak- and mesonotum in lateral view forming slightly convex, Halimun Corridor, Bogor, West Java, 18.ix.2004, (JV04- continuous dorsal outline. Metanotal groove in dorsal SKY-66) (Sk. Yamane) (in soil); one worker, Batang Gadis view transverse, almost straight in median portion, N. P., 1200 m alt., N. Sumatra (secondary forest), forming deep concavity that is laterally margined by 4.v.2006 (Syaukani); one worker, Lubuk Mintrun nr lamellate ridges. Propodeal spiracles oval, situated at Padang, W. Sumatra, 25.vii.1985, (S. & Sk. Yamane); posterolateral corners of propodeum, apart from six workers, Surisura, Pulau Siberut, Mentawai Is., metapleural gland bullae. Propodeal spines devel- W. Sumatra (SU07-SKY-032) (Sk. Yamane); ﬁve workers, oped, longer than diameter of propodeal spiracles, in Sumberjaya (800–900 m alt.), Lampung Barat, Sumatra, dorsal view directed posteriad. 18–19.ix.2007 (nest in soil) (SU07-SKY-213) (Sk. Petiole in dorsal view with subparallel sides and Yamane); two workers, Sumberjaya (800–900 m alt.), narrow anteriorly, longer than wide. Posterior portion Lampung Barat, Sumatra, 18.ix.2007 (coffee planta- of petiole with short process that is slightly higher than tion) (Sk. Yamane); one worker, Batang Gadis N.P. posterior margin of petiole disc in lateral view. (1200 m alt.) (secondary forest), 4.v.2006 (Syaukani); Subpetiolar process developed as blunt process. MALAYSIA: four workers, 19 miles, Cameron High- Postpetiole in lateral view with weakly convex dorsum, lands, 10.iii.2010 (SH10-Mal-24) (S. Hosoishi); six as high as petiole, in dorsal view as wide as or slight- workers, 7 miles, Cameron Highlands, 9.iii.2010 (SH10- ly wider than petiole, weakly bilobed posteriorly but Mal-17) (S. Hosoishi); four workers, Ulu Gombak, without longitudinal sulcus. Subpostpetiolar process Selangor, 3.xii.2005, (SH05-Mal-24) (S. Hosoishi); 15 developed bluntly. workers, UKM Forest, Fraser’s Hill, 15.iii.2005, Integument essentially sculptured. Dorsal surface of (2005c31) (S. Hosoishi); THAILAND: one worker, Khao head generally smooth, but with rugulae on surround- Nam Kang National Park, Songkhla Prov., S. Thai- ing region of antennal sockets. Mandibles with feeble land, 25.vii.1997 (Sk. Yamane); four workers, Khlong rugulae and smooth interspaces. Clypeus generally Klai Stn., Khao N. P., Nakhon S. Thammarat, S. Thai- smooth, but with one distinct pair of longitudinal land, 13.iii.2007 (TH07-SKY-85) (Sk. Yamane). rugulae and one pair of feeble rugulae; distinct rugulae extending to posterior clypeal margin. Anterolateral shoulders of pronotum with rugulae. Lateral surface CREMATOGASTER LUZONENSIS SP. NOV. of pronotum smooth and shining. Pronotum and (FIG. 106) mesonotum with longitudinal rugulae and sculptured interspaces. Mesopleura weakly sculptured. Rugula Holotype worker. Luzon, PHILIPPINES, 25.iv.1992 on higher portion of mesopleura weakly developed. (PH64 coconut) (M. J. Way) (BMNH). Dorsal surface of propodeum generally smooth and Paratypes. One worker, same data as holotype shining, but one pair of rugulae running from metanotal (MPMP); three workers, Luzon, PHILIPPINES, 2.v.1992 groove extending posteriorly and diverging to tips of (PH81 coconut) (M. J. Way) (CASC, KUEC, MHNG). propodeal spines. Dorsal surface of petiole generally smooth. Lateral surface of petiole sculptured. Worker measurements (n = 5): HW 0.41–0.46; HL 0.42– Dorsal and lateral surfaces of postpetiole weakly 0.46; CI 96–100; SL 0.36–0.42; SI 88–95; EL 0.11– sculptured. 0.13; PW 0.26–0.29; WL 0.49–0.55; PSL 0.07–0.1; PtL Standing pilosity sparse. Dorsal face of head with 0.15–0.17; PtW 0.13–0.16; PtH 0.11–0.13; PpL 0.09– some pairs (c. 8) of erect and stout setae, and short 0.11; PpW 0.15–0.17; PtHI 71–81; PtWI 87–94; PpWI and decumbent setae sparsely. Clypeus with two pairs 145–178; WI 100–114. of long and stout setae in anterior portion, one directed upward and the other downward. Anterior clypeal General description of worker: Workers monomorphic. margin with one pair of long setae medially and some Head subquadratic in full-face view. Mandibles with pair of short setae laterally. Scapes with decumbent four teeth arranged at an equal distance, apical and setae. Mesosoma with ﬁve pairs of long erect and stout subapical teeth large, basal two teeth smaller. Ante- setae (ps1PN, psaMN, pspMN, ps1PS, and ps2PS) that rior clypeal margin convex in medial portion. Com- are much longer than other erect setae. Posterolateral

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 586 S. HOSOISHI AND K. OGATA tubercles of petiole posteriorly with one pair of stout portion, forming deep concavity that is laterally mar- setae. Postpetiole with three pairs of stout setae on gined by ridges. Propodeal spiracles oval, situated at disc antrodorsally, anterolaterally and posteriorly. Fourth posterolateral corners of propodeum, apart from abdominal tergite with some pairs (c. > 10) of erect and metapleural gland bullae. Propodeal spines devel- stout setae, and short appressed seate sparsely. oped, distinctly longer than diameter of propodeal spira- Body yellow. All flagellar segments yellow. cles, in dorsal view strongly divergent. Petiole in dorsal view with subparallel sides and Comments: In the worker this species can be distin- narrow short peduncle anteriorly, distinctly longer than guished from all other members of the C. biroi group wide. Posterior portion of petiole with short process by the scape with suberect to decumbent setae, dis- that is slightly higher than posterior margin of petiole tinct compound eyes, generally smooth dorsal surface disc in lateral view. Subpetiolar process developed as of head, and sparse erect and stout setae on body. acute tubercle. Postpetiole in lateral view with strong- This species is similar to C. fritzi, but can be distin- ly convex dorsum, distinctly higher than petiole, in guished from it by the suberect to decumbent dorsal view as wide as petiole, globular, not bilobed. setae on scape, shorter setae on anterior mesonasal Subpostpetiolar process undeveloped, but venter of ridge, abundant erect setae on fourth abdominal postpetiole convex. tergite. Integument essentially sculptured. Dorsal surface of head smooth and shining. Mandibles with rugulae and Distribution: This species is known from the type lo- smooth interspaces. Clypeus generally smooth and cality of the Philippines (Fig. 51). shining, but with one distinct pair of longitudinal rugulae; distinct rugulae not extending to posterior Etymology: The specific name refers to the island of clypeal margin. Anterolateral shoulders of pronotum origin, Luzon. with feeble rugulae. Dorsal surface of pronotum sculptured reticulaterly. Lateral surface of pronotum weakly punctate. Mesopleura generally sculptured. Dorsal CREMATOGASTER MACRACANTHA CREIGHTON, 1945 surface of propodeum sculptured. Dorsal and lateral (FIG. 107) surfaces of petiole sculptured. Dorsal and lateral sur- Crematogaster (Rhachiocrema) macracantha Creighton, faces of postpetiole weakly sculptured. 1945: 114, pl. 12, figs. 4, 5; worker, Mt. Penrissen, Standing pilosity sparse. Dorsal face of head with Sarawak, Borneo, Malaysia (E. Mjoberg) (types not seen). three pairs of long erect setae and appressed setae Crematogaster macracantha; Blaimer, 2012c: 55 [Com- sparsely. Clypeus with two pairs of long setae in ante- bination in C.(Crematogaster)]. rior portion, one directed upward and the other downward. Anterior clypeal margin with one pair of long Worker measurements (n = 3): HW 0.57–0.60; HL 0.61– setae medially and some pairs of short setae lateral- 0.63; CI 93–98; SL 0.65–0.68; SI 111–114; EL 0.14– ly. Scapes with decumbent to appressed setae. Mesosoma 0.15; PW 0.38–0.39; WL 0.84–0.89; PSL 0.50–0.59; PtL with three pairs of long erect and stout setae (ps1PN, 0.37–0.38; PtW 0.21–0.25; PtH 0.19–0.22; PpL 0.18– ps2PN, and psaMN) that are much longer than other 0.20; PpW 0.22–0.24; PtHI 50–58; PtWI 57–66; PpWI erect setae. Posterolateral tubercles of petiole poste- 118–122; WI 88–105. riorly with one pair of long setae. Postpetiole with three pairs of long setae on disc anteriorly, posteriorly and General description of worker: Workers monomorphic. laterally. Fourth abdominal tergite with erect setae Head round in full-face view. Mandibles with four sparsely, but no decumbent to appressed setae. teeth arranged at an equal distance, apical and Body yellow-brown. Apical two flagellar segments light subapical teeth large, basal two teeth smaller. Ante- yellow, contracting with other flagellar segments that rior clypeal margin convex in medial portion. Com- are yelow-brown. pound eyes distinctly projecting beyond lateral margins of head in full-face view. Scapes exceeding posterolateral Comments: We have not been able to examine the types corners of head. of C. macracantha, but the original description and Pronotal collar with concave anterior margin in dorsal figures closely match the specimens examined. In the view, slightly lower than pronotum in lateral view. worker this species can be easily distinguished from Pronotal dorsum without distinct longitudinal ridges the other species of the C. baduvi group by the long laterally. Mesonotal dorsum with lateral ridges that propodeal spines (PSL 0.50–0.59 vs. 0.15–0.22 in the irregularly extend posteriad to tips of propodeal spines. others) and sculptured surface of the body. Pronotum and mesonotum in lateral view forming evenly Blaimer (2012c) assigned C. macracantha in the sub- arched, continuous dorsal outline. Metanotal groove in genus Crematogaster in the subgeneric revision, but dorsal view transverse, almost straight in median she did not include the species in molecular phylogenetic

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 587 analysis (2012a, 2012c). Although we do not have any view transverse, almost straight in median portion, sequence data for the species, several morphological forming deep concavity that is laterally margined by features: clearly differentiated two-segmented antennal thin lamellate ridges. Propodeal spiracles oval, situ- club; laterally directed propodeal spines; petiole with ated at posterolateral corners of propodeum, touch- subparallel sides; node-like process on posterior portion ing metapleural gland bullae. Propodeal spines of petiole; highly convex postpetiolar dorsum; developed, longer than diameter of propodeal spira- subpostpetiolar venter convex, suggest that C. cles, in dorsal view directed posteriad. macracantha belongs to the subgenus Orthocrema. Thus Petiole in dorsal view with subparallel sides or weakly we treat C. macracantha as a member of the subge- convex sides and narrow anteriorly, longer than wide. nus Orthocrema. Posterior portion of petiole without distinct process in lateral view. Subpetiolar process developed, acute api- Distribution: This species is known from Malaysia cally. Postpetiole in lateral view with weakly convex (Borneo) (Fig. 49). dorsum, as high as petiole, in dorsal view as wide as or slightly narrower than petiole, weakly bilobed pos- Material examined: MALAYSIA: one worker, Bt. teriorly, but without longitudinal sulcus. Subpostpetiolar Entimau (390 m alt.), Sarawak, Borneo, 18.iv.1994, process developed as small process. (Mahmud) (P1B4); two workers, Poring (600–700 m alt.), Integument weakly sculptured. Dorsal surface of head Sabah, Borneo, E. Malaysia, 8.i.1998, (F. Yamane). generally smooth, but with rugulae on surrounding region of antennal sockets. Mandibles with feeble rugulae and smooth interspaces. Clypeus smooth and CREMATOGASTER MASUKOI HOSOISHI,YAMANE & shining without distinct longitudinal rugulae. Dorsal OGATA, 2010 and lateral surfaces of pronotum smooth and shining; (FIG. 108) anterolateral shoulders of pronotum without rugulae. Crematogaster (Orthocrema) masukoi Hosoishi, Yamane Mesopleura smooth on central areas, but sculptured & Ogata, 2010: 347, fig. 1, 3, 5; Holotype worker, with rugulae on their marginal areas. Rugula on higher Sepilok, Sandakan, Borneo, Malaysia, 17.viii.1981 (K. portion of mesopleura developed. One pair of rugulae Masuko) (ITBC, examined). Two paratype workers, same running from metanotal groove extending posteriorly data as holotype (KUEC, MHNG, examined) and five and diverging to propodeal spines. Horizontal rugula paratype workers, Sangkimah, Kutai N. P., E. on dorsal surface of propodeum interruptedly extend- Kalimantan, Indonesia, 13.ix.1993 (P-2 soil-2) ing to tips of propodeal spines. Dorsal surface of (Sk. Yamane) (BMNH, MCZC, MBBJ, KUEC, SKYC, propodeum sculptured. Lateral surface of propodeum examined). generally smooth and shining, but weakly sculptured Crematogaster masukoi; Blaimer, 2012c: 55 [Com- with rugulae on anterior areas. Dorsal surface of petiole bination in C.(Orthocrema)]. generally smooth. Lateral surface of petiole weakly sculptured. Dorsal surface of postpetiole smooth and Worker measurements (n = 6): HW 0.46–0.50; HL 0.44– shining. Lateral surface of postpetiole weakly sculptured. 0.48; CI 100–109; SL 0.35–0.37; SI 74–78; EL 0.04– Standing pilosity sparse. Dorsal face of head with 0.06; PW 0.28–0.31; WL 0.53–0.57; PSL 0.08–0.11; PtL three pairs of long erect and stout setae, and short 0.16–0.18; PtW 0.16–0.17; PtH 0.11–0.13; PpL 0.10– and appressed setae sparsely. Clypeus with two pairs 0.12; PpW 0.15–0.16; PtHI 61–76; PtWI 94–106; PpWI of long and stout setae in anterior portion, one direct- 133–160; WI 88–100. ed upward and the other downward. Anterior clypeal margin with one single short setae medially and one General description of worker: Workers monomorphic. pair of long setae laterally, and some pair of short setae Head subquadratic in full-face view. Mandibles with laterally. Scapes with suberect to decumbent setae. four teeth, basal tooth arranged away from the third Mesosoma with five pairs of long erect and stout setae apical one, apical and subapical teeth large, basal two (ps1PN, psaMN, pspMN, ps1PS, and ps2PS) that are teeth smaller. Anterior clypeal margin weakly convex much longer than other erect setae. Posterolateral tu- or almost straight in medial portion. Compound eyes bercles of petiole posteriorly with two pairs of stout not projecting beyond lateral margins of head in full- setae. Postpetiole with three pairs of stout setae on face view. Scapes reaching posterolateral corners of head. disc anterodorsally, anterolaterally, posteriorly. Fourth Pronotal collar with almost straight anterior margin abdominal tergite with suberect setae sparsely, but no in dorsal view, distinctly lower than pronotum in lateral decumbent to appressed setae. view. Pronotal dorsum without distinct ridges later- Body yellow to brown. All flagellar segments yellow. ally. Mesonotal dorsum without lateral ridges. Pronotum and mesonotum in lateral view forming slightly convex, Comments: In the worker this species is very distinct continuous dorsal outline. Metanotal groove in dorsal among the C. biroi group in having reduced com-

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 588 S. HOSOISHI AND K. OGATA pound eyes (with c. 6 ommatidia). It is similar to C. Subpetiolar process undeveloped. Postpetiole in lateral myops (C. quadriruga group) in having reduced com- view with weakly convex dorsum, as high as petiole, pound eyes, but can be easily distinguished from it by in dorsal view as wide as petiole, globular, not bilobed. the smooth surface of clypeus, developed subpetiolar Subpostpetiolar process undeveloped. process and acutely developed subpostpetiolar process. Integument essentially smooth and shining. Dorsal surface of head smooth and shining. Gena with lon- Distribution and biology: This species is known from gitudinal rugulae. Mandibles with feeble rugulae and Malaysia (Borneo) and Indonesia (Kalimantan) (Fig. 51). smooth interspaces. Clypeus generally smooth and This species inhabits developed forests, and nests in shining, but with one to two distinct pairs of longitu- soil. dinal rugulae; longer rugulae extending to posterior clypeal margin. Dorsal and lateral surfaces of pronotum Material examined: INDONESIA: four workers, smooth and shining; anterolateral shoulders of pronotum Sangkimah, Kutai N. P., E. Kalimantan, 13.ix.1993 (P-2 without rugulae. Mesopleura generally smooth and soil-2) (Sk. Yamane). shining. Rugula on higher portion of mesopleura extending to small pit of mesothoracic spiracles. Dorsal surface of propodeum generally smooth and shining. CREMATOGASTER MOATENSIS SP. NOV. Dorsal surface of petiole smooth and shining. Lateral (FIG. 109) surface of petiole generally smooth, without longitu- Holotype worker. nr. Kotamobaqu (1200 m), Danau Moat, dinal rugulae. Dorsal and lateral surfaces of postpetiole Utara, Sulawesi, INDONESIA, 11.ix.1985 (no collec- smooth and shining. tor’s name) (BMNH). Standing pilosity sparse. Dorsal face of head with Paratype. Five workers, same data as holotype erect to suberect setae sparsely. Clypeus with three (BMNH, CASC, KUEC, MBBJ, THNHM). pairs of long setae in anterior portion, one directed upward, one downward, the other laterally below antennal Worker measurements (n = 6): HW 0.46–0.49; HL 0.48– sockets. Anterior clypeal margin with single long setae 0.53; CI 92–98; SL 0.45–0.47; SI 94–100; EL 0.11– medially and one pair of long setae laterally, and short 0.12; PW 0.30–0.33; WL 0.60–0.64; PSL 0.12–0.13; PtL setae laterally. Gena (malar space) with some suberect 0.19–0.20; PtW 0.14–0.15; PtH 0.13–0.14; PpL 0.10– setae near mandibular insertion. Scapes with suberect 0.11; PpW 0.15–0.16; PtHI 68–70; PtWI 70–79; PpWI to decumbent setae. Mesosoma with four pairs of long 136–160; WI 100–114. erect and stout setae (ps1PN, psaMN, pspMN, and ps1PS/ ps2PS) that are much longer than other erect setae. General description of worker: Workers monomorphic. Posterolateral tubercles of petiole posteriorly with one Head subquadratic in full-face view. Mandibles with pair of long setae. Postpetiole with three pairs of long four teeth arranged at an equal distance, apical and setae on disc anterodorsally, anterolaterally, and pos- subapical teeth large, basal two teeth smaller. Ante- teriorly. Fourth abdominal tergite with erect to suberect rior clypeal margin convex in medial portion. Com- setae sparsely, but no decumbent to appressed setae. pound eyes distinctly projecting beyond lateral margins Body bicolored, head and gaste with brown, mesosoma of head in full-face view. Scapes reaching posterolateral with yellow. All flagellar segments yellow. corners of head. Pronotal collar with weakly concave anterior margin Comments: In the worker this species is very unique in dorsal view, distinctly lower than pronotum in lateral among the Asian Orthocrema fauna in having the view. Pronotal dorsum without ridges laterally. propodeal spines directed posteriad and the metanotal Mesonotal dorsum without distinct lateral ridges pos- groove laterally not defined by lamellate ridges. teriorly. Pronotum and mesonotum in lateral view forming slightly convex, continuous dorsal outline. Metanotal groove in dorsal view transverse, almost Distribution: This species is known only from Indo- straight in median portion, forming deep concavity that nesia (Sulawesi) (Fig. 49). is laterally margined by ridges. Propodeal spiracles oval, situated at posterolateral corners of propodeum, apart Etymology: The specific name refers to the type from metapleural gland bullae. Propodeal spines de- locality, Danau Moat (Lake Moat). veloped, longer than diameter of propodeal spiracles, in dorsal view directed posteriad. Petiole in dorsal view with subparallel sides and Material examined: INDONESIA: three workers, narrow anteriorly, longer than wide. Posterior portion Dumoga-Bone N. P., Utara, Sulawesi, 17.iv.1985 (‘Clarke’ of petiole with short process that is slightly higher than Camp) (lower montane forest, 1140 m) (no collector’s posterior margin of petiole disc in lateral view. name).

CREMATOGASTER MYOPS FOREL, 1911a rugulae on surrounding region of antennal sockets. Man- (FIG. 110) dibles with feeble rugulae and smooth interspaces. Clypeus weakly sculptured with one distinct pair of Crematogaster myops Forel, 1911a: 31; Lectotype and longitudinal rugulae; rugulae not extending to pos- ﬁve paralectotype workers, Sarawak, Borneo, Malay- terior clypeal margin. Dorsal and lateral surfaces of sia (Haviland) (MHNG, examined). pronotum smooth and shining; anterolateral shoul- Crematogaster myops; Santschi, 1918: 182 [Combi- ders of pronotum without rugulae. Mesopleura smooth nation in C.(Orthocrema)]. and shining. Dorsal surface of propodeum generally Crematogaster myops; Emery, 1922: 132 [Combina- smooth and shining, but one pair of rugulae running tion in C.(Orthocrema)]. from metanotal groove to tips of propodeal spines. Dorsal Crematogaster myops; Hosoishi et al. 2010: 347 surface of petiole smooth and shining. Lateral surface [Lectotype designation and redescription of type of petiole generally smooth, but with one longitudi- material]. nal rugula. Dorsal and lateral surfaces of postpetiole Crematogaster myops; Blaimer, 2012c: 55 [Combi- smooth and shining. nation in C.(Orthocrema)]. Standing pilosity sparse. Dorsal face of head with suberect setae sparsely. Clypeus with two pairs of long Worker measurements (n = 7): HW 0.44–0.48; HL 0.45– setae in anterior portion, one directed upward and 0.52; CI 92–98; SL 0.40–0.44; SI 90–94; EL 0.05– the other downward. Anterior clypeal margin with 0.06; PW 0.27–0.31; WL 0.49–0.58; PSL 0.05–0.08; PtL one pair of long setae medially and short setae later- 0.15–0.21; PtW 0.13–0.16; PtH 0.10–0.12; PpL 0.09– ally. Scapes with suberect setae. Mesosoma with four 0.13; PpW 0.12–0.16; PtHI 55–67; PtWI 75–89; PpWI pairs of long erect and stout setae (ps1PN, psaMN, 115–150; WI 92–100. pspMN, and ps1PS) that are much longer than other erect setae. Posterolateral tubercles of petiole poste- General description of worker: Workers monomorphic. riorly with one pair of stout setae. Postpetiole with Head subquadratic in full-face view. Mandibles with several pairs of long setae on disc anteriorly, posteri- four teeth, basal tooth arranged away from the third orly and laterally. Fourth abdominal tergite with apical one, apical and subapical teeth large, basal two suberect setae abundantly, and short decumbent setae teeth smaller. Anterior clypeal margin convex in medial sparsely. portion. Compound eyes not projecting beyond lateral Body yellow. All ﬂagellar segments yellow. margins of head in full-face view. Scapes reaching posterolateral corners of head. Pronotal collar with weakly concave anterior margin Comments: In the worker this species is very distinct in dorsal view, distinctly lower than pronotum in lateral among the C. quadriruga group in having reduced com- view. Pronotal dorsum without distinct ridges later- pound eyes (with c. 6 ommatidia). It is similar to C. ally. Mesonotal dorsum with lateral ridges posteri- masukoi (C. biroi group) in having reduced com- orly. Pronotum and mesonotum in lateral view forming pound eyes, but can be distinguished by the clypeus slightly convex, continuous dorsal outline. Metanotal with rugulae, smooth surface of propodeum, and the groove in dorsal view transverse, almost straight in lack of subpostpetiolar process. median portion, forming deep concavity that is laterally margined by lamellate ridges. Propodeal spira- Distribution and biology: This species is known from cles oval, situated at posterolateral corners of Malaysia (Peninsula and Borneo), Singapore and In- propodeum, touching to metapleural gland bullae. donesia (Sumatra) (Fig. 52). This species inhabits de- Propodeal spines developed, as long as diameter of veloped forests, and nests in soil. propodeal spiracles, in dorsal view directed posteriad. Petiole in dorsal view with subparallel sides and narrow anteriorly, longer than wide. Posterior portion Material examined: INDONESIA: three workers, Bt. of petiole with short process that is slightly higher than Lawang Lowland G. Leuser N. P., N. Sumatra, 17.viii.2002 posterior margin of petiole disc in lateral view. (Sk. Yamane); MALAYSIA: one worker, Tower Region, Subpetiolar process developed as small acute process. Lambir N. P., Miri, Sarawak, 15.i.1993, (Sk. Yamane) Postpetiole in lateral view with weakly convex dorsum, (Canopy Ecol.); one worker, Bako Nat. Park, Sarawak, as high as petiole, in dorsal view as wide as petiole, Borneo, 21–22.iv.1993, (Sk. Yamane); one worker, Danum weakly bilobed posteriorly but without longitudinal Balley, Sabah, Borneo, E. Malaysia, 3–4.iii.1999, sulcus. Subpostpetiolar process undeveloped, but venter (Sk. Yamane); four workers, Old Tower R., Lambir of postpetiole convex. National Park, Sarawak, Borneo, 30.vi.2004 (Sk. Integument essentially smooth and shining. Dorsal Yamane); SINGAPORE: one worker, 4.xii.1995, (Sk. surface of head generally smooth and shining, but with Yamane).

CREMATOGASTER OCELLATA SP. NOV. surface of pronotum generally smooth and shining, (FIG. 111) but with longitudinal rugulae on higher portion extending to mesopleura and lateral sides of propodeum. Holotype worker. Tu Lung, Mai Chau Dist., Hoa Binh Mesopleura sculptured. Rugula on higher portion Prov., VIETNAM, S 20 44 E 104 56, 27.xi.1999 (15 min of mesopleura extending to small pit of mesothoracic TUS) (K. Ogata) (IEBR). spiracles. One pair of rugulae running from metanotal Paratypes. Three workers same data as holotype groove to tips of propodeal spines. Dorsal surface of (CASC, KUEC, THNHM). propodeum weakly sculptured. Dorsal surface of petiole smooth and shining. Lateral surface of petiole sculp- Worker measurements (n = 4): HW 0.50–0.56; HL 0.53– tured. Dorsal surface of postpetiole smooth and shining. 0.59; CI 94–96; SL 0.40–0.45; SI 78–84; EL 0.07– Lateral surface of postpetiole weakly sculptured 0.08; PW 0.31–0.37; WL 0.58–0.66; PSL 0.08–0.10; PtL posteriorly. 0.18–0.22; PtW 0.16–0.18; PtH 0.12–0.15; PpL 0.12– Standing pilosity sparse. Dorsal face of head with 0.13; PpW 0.17–0.18; PtHI 63–78; PtWI 81–85; PpWI erect to suberect and stout long setae sparsely. Clypeus 138–142; WI 100–106. with two pairs of long setae in anterior portion, one directed upward and the other downward. Anterior General description of woker: Workers monomorphic. clypeal margin with one pair of long setae medially Head subquadratic in full-face view. Mandibles with and some pairs of short setae laterally. Scapes with four teeth arranged at an equal distance, apical and suberect setae. Mesosoma with five pairs of long erect subapical teeth large, basal two teeth smaller. Ante- and stout setae (ps1PN, ps2PN, psaMN, pspMN, and rior clypeal margin convex in medial portion. Com- ps1PS) that are much longer than other erect setae. pound eyes not projecting beyond lateral margins of Posterolateral tubercles of petiole posteriorly with two head in full-face view. Scapes reaching posterolateral pairs of stout long setae. Postpetiole with five pairs corners of head. of long setae on disc anterodorsally, anterolaterally and Pronotal collar with almost straight anterior margin posteriorly. Fourth abdominal tergite with erect and in dorsal view, distinctly lower than pronotum in lateral stout suberect long setae sparsely, and short decum- view. Pronotal dorsum without ridges laterally. bent setae sparsely. Mesonotal dorsum with lateral ridges that irregular- Body yellow. All flagellar segments yellow. ly extend posteriad to tips of propodeal spines. Pronotum and mesonotum in lateral view forming slightly convex, Comments: In the worker this species is distinct among continuous dorsal outline. Metanotal groove in dorsal the C. biroi group in having reduced compound eyes view transverse, almost straight in median portion, (with c. 12–15 ommatidia). forming deep concavity that is laterally margined by This species corresponds to sp. eg-12 by Eguchi, Bui lamellate ridges. Propodeal spiracles oval, situated at & Yamane (2011). posterolateral corners of propodeum, touching metapleural gland bullae. Propodeal spines devel- Distribution: This species is known from N. Vietnam oped, longer than diameter of propodeal spiracles, in (Fig. 51). dorsal view directed posteriad. Petiole in dorsal view with subparallel sides and Etymology: The specific name refers to the small com- narrow anteriorly, longer than wide. Posterior portion pound eyes. of petiole with short process that is slightly higher than posterior margin of petiole disc in lateral view. Material examined: VIETNAM: one worker, Liem Phu Subpetiolar process weakly developed as blunt process. (450–600 m alt.), Van Ban Dist., Lao Cai Prov., 27 Postpetiole in lateral view with weakly convex dorsum, ix.2006 (K. Eguchi). as high as petiole, in dorsal view as wide as petiole, globular, not bilobed. Subpostpetiolar process devel- CREMATOGASTER OSAKENSIS FOREL, 1900 oped as blunt process. Integument essentially smooth and shining. Dorsal (FIG. 112) surface of head generally smooth and shining, but Crematogaster sordidula var. osakensis Forel, 1900: 269; weakly with rugulae on surrounding region of antennal syntype workers, Osaka, Japan (MHNG, examined) sockets. Mandibles with feeble rugulae and smooth (Synonymy under C. japonica by Brown, 1949: 37). interspaces. Clypeus generally smooth and shining, One syntype worker in MHNG here designated but with one distinct pair of longitudinal rugulae; Lectotype. rugulae not extending to posterior clypeal margin. Crematogaster sordidula var. japonica Forel, 1912c: Anterolateral shoulders of pronotum with feeble rugulae. 339; syntype workers, Tokyo, Japan (Ito) (MHNG, ex- Dorsal surface of pronotum smooth and shining. Lateral amined) (Synonymy under C. osakensis, by Brown, 1949:

Figures 112–119. Body in lateral view. 112, Crematogaster osakensis, worker [Hiraodai Karst Plateau, Fukuoka, Japan]; 113, Crematogaster philippinensis, worker [Mt. Mayon, Luzon Island, Philippines]; 114, Crematogaster quadriruga, worker [Lum Jang Wat, E, Thailand]; 115, Crematogaster quadriruga, intermediate worker [PSU forest, Hat Yai, Songkhla Prov., S. Thailand]; 116, Crematogaster reticulata, worker [Ulu Gombak, Selangor, W. Malaysia]; 117, Crematogaster reticulata, intermediate worker [Datfa Waterfall, Tai Rom Yen National Park, Surat Thani Prov., S. Thailand]; 118, Crematogaster schimmeri, worker [Pilam, Taiwan]; 119, Crematogaster storki, worker [Dumoga-Bone National Park, Utara, Sulawesi, Indonesia].

37). One syntype worker (top specimen of three on one dorsum, as high as petiole, in dorsal view as wide as pin) in MHNG here designated Lectotype. petiole, globular, not bilobed. Subpostpetiolar process Crematogaster sordidula subsp. osakensis; Emery, developed as blunt process. 1912: 671 [Subspecies of sordidula]. Integument essentially smooth and shining. Dorsal Crematogaster sordidula subsp. osakensis; Emery, surface of head generally smooth and shining, but with 1922: 131 [Combination in C.(Orthocrema)]. rugulae on surrounding region of antennal sockets. Man- Crematogaster sordidula var. japonica; Emery, 1922: dibles with feeble rugulae and smooth interspaces. 131 [Combination in C.(Orthocrema)]. Clypeus generally smooth and shining, but with two Crematogaster sordidula subsp. osakensis; Wheeler, distinct pairs of longitudinal rugulae; rugulae not ex- 1928: 111 [Subspecies of sordidula; descriptions of queen tending to posterior clypeal margin. Anterolateral shoul- and male]. ders of pronotum with rugulae. Dorsal surface of Crematogaster osakensis; Collingwood, 1976: 303 pronotum with rugulae. Lateral surface of pronotum [Raised to species]. smooth and shining. Mesopleura weakly sculptured, Crematogaster osakensis; Onoyama, 1980: 198 [Sub- but relatively smooth on central areas. Rugula on higher species of sordidula]. portion of mesopleura extending to small pit of Crematogaster osakensis; Kupyanskaya, 1990: 129 mesothoracic spiracles. Dorsal surface of propodeum [Raised to species]. with reticulated rugulae. Dorsal surface of petiole smooth Crematogaster osakensis; Bolton, 1995: 159. and shining. Lateral surface of petiole weakly sculp- Crematogaster osakensis; Blaimer, 2012c: 55 [Com- tured. Dorsal surface of postpetiole smooth and shining. bination in C.(Orthocrema)]. Lateral surface of postpetiole weakly sculptured posteriorly. Worker measurements (n = 8): HW 0.47–0.57; HL 0.49– Standing pilosity sparse. Dorsal face of head with 0.59; CI 92–100; SL 0.38–0.44; SI 77–84; EL 0.11– three pairs of erect and stout long setae, and short 0.13; PW 0.27–0.35; WL 0.54–0.68; PSL 0.08–0.11; PtL and appressed setae sparsely. Clypeus with two pairs 0.16–0.19; PtW 0.15–0.18; PtH 0.12–0.15; PpL 0.11– of long setae in anterior portion, one directed upward 0.12; PpW 0.14–0.18; PtHI 68–81; PtWI 83–94; PpWI and the other downward. Anterior clypeal margin with 117–150; WI 88–100. one single long setae medially and one pairs of long setae laterally, and some pairs (three to four) of short General description of worker: Workers monomorphic. setae laterally. Scapes with suberect to decumbent setae. Head subquadratic in full-face view. Mandibles with Mesosoma with three pairs of long erect and stout setae four teeth arranged at an equal distance, apical and (ps1PN, psaMN, and pspMN) that are much longer than subapical teeth large, basal two teeth smaller. Ante- other setae and one pair of shorter setae (ps1PN). rior clypeal margin slightly concave in medial portion. Posterolateral tubercles of petiole posteriorly with one Compound eyes distinctly projecting beyond lateral pair of stout long setae. Postpetiole with three pairs margins of head in full-face view. Scapes reaching of stout long setae on disc anterodorsally, anterolaterally posterolateral corners of head. and posteriorly. Fourth abdominal tergite with suberect Pronotal collar with almost straight anterior margin to decumbent stout setae sparsely. in dorsal view, distinctly lower than pronotum in lateral Body yellow. All ﬂagellar segments yellow. view. Pronotal dorsum with ridges laterally. Mesonotal dorsum with lateral ridges posteriorly that irregular- Comments: In the worker this species can be distin- ly extend posteriad to tips of propodeal spines. Pronotum guished from all other members of the C. biroi group and mesonotum in lateral view forming convex, con- by the distinct compound eyes, generally smooth dorsal tinuous dorsal outline. Metanotal groove in dorsal surface of head, petiole tapering posteriorly in dorsal view transverse, almost straight in median portion, view, and erect setae on body tapering distally. This forming deep concavity that is laterally margined by species is similar to C. vieti, but can be distinguished lamellate ridges. Propodeal spiracles oval, situated at from it by the slender propodeal spines, petiole taper- posterolateral corners of propodeum, touching ing posteriorly and subpostpetiolar process angulate. metapleural gland bullae. Propodeal spines developed, longer than diameter of propodeal spiracles, in Distribution and biology: This species is known from dorsal view directed posteriad. Japan (Hokkaido, Honshu, Shikoku, Kyushu, Yakushima Petiole in dorsal view with subparallel sides and Island, Amami Island) (Japanese Ant Database Group weakly angulate shoulders anteriorly, longer than wide. 2008, 2008), South Korea (Terayama, Choi & Kim, 1992; Posterior portion of petiole with short process that is Choi, Ogata & Terayama, 1993), North Korea slightly higher than posterior margin of petiole disc (Collingwood, 1976; Radchenko, 2005) and China (Wu in lateral view. Subpetiolar process developed as acute & Wang, 1995; Zhou, 2001) (Fig. 51). It is noted that process. Postpetiole in lateral view with weakly convex we have not examined the specimens from South Korea,

North Korea and China in this study. This species in- Petiole in dorsal view with subparallel sides and habits grasslands (Hosoishi et al., 2015) to forests, and narrow anteriorly, longer than wide. Posterior portion nests under stone and in soil or leaf litter. Colonies of petiole with short process that is slightly higher than are polygynous. Reproductive alates fly at evening in posterior margin of petiole disc in lateral view. September and are attracted to light (Japanese Ant Subpetiolar process undeveloped. Postpetiole in lateral Database Group 2008, 2008). A myrmecophilous beetle, view with weakly convex dorsum, as high as petiole, Triartiger reductus Nomura (Staphylinidae) are known in dorsal view as wide as petiole, weakly bilobed pos- from the nests in Tsushima Island, Japan (Komatsu teriorly but without longitudinal sulcus. Subpostpetiolar & Maruyama, 2008). process undeveloped, but venter of postpetiole convex. Integument essentially smooth and shining. Dorsal Material examined: JAPAN: four workers, Takuhi surface of head smooth and shining. Mandibles with Shrine, Nishinoshima, Oki Islands, Shimane, 17.ix.2003 feeble rugulae and smooth interspaces. Clypeus gen- (T. Yamauchi); 30 workers, Hiraodai Karst Plateau, erally smooth and shining, but with one pair of lon- Fukuoka, 28.v.2010 (S. Hosoishi); six workers, Nomozaki, gitudinal rugulae; rugulae not extending to posterior Nagasaki, 27.vii.1978 (K. Ogata); 11 workers, Anbou, clypeal margin. Dorsal and lateral surfaces of pronotum Yakushima Island, Kagoshima, 25.viii.2004 (K. Ogata smooth shining; anterolateral shoulders of pronotum et al.). with rugulae. Mesopleura smooth and shining. Dorsal surface of propodeum generally smooth and shining, but one pair of rugulae running from metanotal groove CREMATOGASTER PHILIPPINENSIS SP. NOV. to tips of propodeal spines. Dorsal surface of petiole (FIG. 113) smooth and shining. Lateral surface of petiole Holotype worker. Mt. Makiling, Los Banos, Luzon Island, generally smooth, but with one longitudinal rugula. PHILIPPINES, 13.vii.1997 (Sk. Yamane) (MPMP). Dorsal and lateral surfaces of postpetiole smooth and Paratype. Four workers, same data as holotype shining. (BMNH, CASC, KUEC, SKYC). Standing pilosity sparse. Dorsal face of head with suberect setae sparsely. Clypeus with two pairs of long Worker measurements (n = 5): HW 0.45–0.46; HL 0.48– setae in anterior portion, one directed upward and the 0.49; CI 94–96; SL 0.41–0.43; SI 91–93; EL 0.09– other downward. Anterior clypeal margin with one pair 0.11; PW 0.28–0.31; WL 0.56–0.59; PSL 0.08–0.11; PtL of long setae medially and short setae laterally. Scapes 0.16–0.18; PtW 0.13–0.14; PtH 0.13–0.14; PpL 0.11– with suberect setae. Mesosoma with four pairs of long 0.12; PpW 0.14–0.15; PtHI 72–82; PtWI 76–88; PpWI erect and stout setae [ps1PN (two pairs), psaMN, 117–127; WI 100–108. pspMN, and ps1PS] that are much longer than other erect setae. Posterolateral tubercles of petiole poste- General description of worker: Workers monomorphic. riorly with one pair of stout setae (one pair of short Head subquadratic in full-face view. Mandibles with setae laterally). Postpetiole with three pairs of setae four teeth arranged at an equal distance, apical and on disc anteriorly, posteriorly and laterally. Fourth ab- subapical teeth large, basal two teeth smaller. Ante- dominal tergite with erect to suberect setae abundant- rior clypeal margin convex in medial portion. Com- ly, and short decumbent setae sparsely. pound eyes distinctly projecting beyond lateral margins Body yellow. All flagellar segments yellow. of head in full-face view. Scapes reaching posterolateral corners of head. Comments: In the worker this species can be distin- Pronotal collar with weakly concave anterior margin guished from all other members of the C. quadriruga in dorsal view, distinctly lower than pronotum in lateral group by the distinct compound eyes, V-shaped view. Pronotal dorsum without distinct ridges later- metanotal groove in lateral view, large propodeal spira- ally. Mesonotal dorsum with lateral ridges posteri- cles touching metapleural gland bulla, long propodeal orly that irregularly extend posteriad to tips of propodeal spines (PSL 0.08–0.11), and yellow-colored body. This spines. Pronotum and mesonotum in lateral view not species is similar to C. sundalandensis, but can be dis- clearly forming continuous dorsal outline; mesonotal tinguished from it by the scape with suberect setae dorsum flat. Metanotal groove in dorsal view trans- only and yellow-colored body. verse, almost straight in median portion, forming deep concavity that is laterally margined by lamellate ridges. Propodeal spiracles elliptical, situated at posterolateral Distribution: This species is known from the type lo- corners of propodeum, touching to metapleural gland cality of the Philippines (Fig. 52). bullae. Propodeal spines developed, longer than diameter of propodeal spiracles, in dorsal view directed Etymology: The specific name refers to the country of posteriad. origin, the Philippines.

Material examined: PHILIPPINES: four workers, Mt. Integument essentially smooth and shining. Dorsal Mayon, Luzon Island, 28.xi.2005 (T. Kobayashi); six surface of head smooth and shining. Mandibles with workers, Oriental Valencia, Negros, 25.v.1983 (676) (C. feeble rugulae and smooth interspaces. Clypeus gen- K. Stall & F. P. Godoy). erally smooth and shining, but with one distinct pair of longitudinal longer rugulae and one pair of shorter rugulae laterally; longer rugulae not extending to pos- CREMATOGASTER QUADRIRUGA FOREL, 1911c terior clypeal margin. Dorsal and lateral surfaces of STAT. NOV. pronotum smooth and shining; anterolateral shoul- (FIGS 114, 115) ders of pronotum with longitudinal rugulae. Mesopleura Crematogaster biroi var. quadriruga Forel, 1911c: 455; weakly punctuate, but seems smooth and shining. Dorsal syntype workers, Kerr, Thailand (M. Waldo) (MHNG, surface of propodeum generally smooth and shining, examined). One syntype worker (top specimen of three but with longitudinal rugulae anteriorly. Dorsal on one pin) in MHNG here designated Lectotype. and lateral surfaces of petiole smooth and shining. Crematogaster biroi var. quadriruga; Emery, 1922: Dorsal and lateral surfaces of postpetiole smooth and 132 [Combination in C.(Orthocrema)]. shining. Standing pilosity sparse. Dorsal face of head with Worker measurements (n = 8): HW 0.46–0.50; HL 0.48– erect setae sparsely. Clypeus with two pairs of long 0.51; CI 94–100; SL 0.44–0.48; SI 94–100; EL 0.11– setae in anterior portion, one directed upward and the 0.12; PW 0.27–0.31; WL 0.54–0.60; PSL 0.11–0.13; PtL other downward. Anterior clypeal margin with one pair 0.17–0.22; PtW 0.14–0.17; PtH 0.13–0.15; PpL 0.12– of long setae medially and short setae laterally. Scapes 0.17; PpW 0.15–0.18; PtHI 68–79; PtWI 74–85; PpWI with suberect and decumbent setae. Mesosoma with 100–145; WI 100–113. five to six pairs of long erect and stout setae (ps1PN, psaMN, pspMN, ps1PS, and one to two ps2PS) that General description of worker: Workers monomorphic, are much longer than other erect setae. Posterolateral but intermediate worker as large as queen. tubercles of petiole posteriorly with one pair of stout Head subquadratic in full-face view. Mandibles with long setae and one pair of short setae laterally. four teeth arranged at an equal distance, apical and Postpetiole with three pairs of setae on disc subapical teeth large, basal two teeth smaller. anterodorsally, anterolaterally and posteriorly. Fourth Anterior clypeal margin convex in medial portion. abdominal tergite with erect setae abundantly, and short Compound eyes distinctly projecting beyond lateral appressed setae sparsely. margins of head in full-face view. Scapes reaching Body bicolored with head, mesosoma, petiole, posterolateral corners of head. postpetiole and first gater yellow and with the remains Pronotal collar with weakly concave anterior margin of the gaster brown. All flagellar segments yellow. in dorsal view, distinctly lower than pronotum in lateral view. Pronotal dorsum with ridges laterally. Mesonotal Intermediate worker measurements (n = 2): HW 0.65– dorsum with lateral ridges that irregularly extend 0.68; HL 0.66–0.68; CI 98–100; SL 0.60–0.61; SI 90– posteriad to tips of propodeal spines. Pronotum and 92; EL 0.17; PW 0.48–0.51; WL 0.89–0.92; PSL 0.15; mesonotum in lateral view not clearly forming con- PtL 0.35; PtW 0.25–0.26; PtH 0.23–0.25; PpL 0.21– tinuous dorsal outline; mesonotal dorsum higher an- 0.22; PpW 0.28–0.29; PtHI 66–71; PtWI 71–74; PpWI teriorly. Metanotal groove in dorsal view transverse, 131–133; WI 111–112. almost straight in median portion, forming deep concavity that is laterally margined by lamellate ridges. Description of intermediate worker: With worker char- Propodeal spiracles oval, situated at posterolateral corners acter conditions, except as follows. of propodeum, apart from metapleural gland bullae. Three ocelli present. Propodeal spines developed, longer than diameter of Mesonotum strongly convex in lateral view. Posteri- propodeal spiracles, in dorsal view directed posteriad. or face of mesonotum steeply sloping so that in lateral Petiole in dorsal view with subparallel sides and view dorsal outline of promesonotum not smoothly narrow anteriorly, longer than wide. Posterior portion arched. of petiole with short process that is slightly higher than Subpetiolar process developed as blunt process. posterior margin of petiole disc in lateral view. Dorsal surface of pronotum generally smooth, but Subpetiolar process developed as acute process. with feeble rugulae. Dorsal surface of propodeum weakly Postpetiole in lateral view with weakly convex dorsum, punctuated with longitudinal rugulae. Lateral surface as high as petiole, in dorsal view as wide as petiole, of petiole weakly sculptured. Dorsal surface of postpetiole weakly bilobed posteriorly but without longitudinal smooth and shining. sulcus. Subpostpetiolar process undeveloped, but venter Posterolateral tubercles of petiole with three to of postpetiole convex. four pais of stout long setae posteriorly. Fourth

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 595 abdominal tergite with erect setae abundantly, and short Kr. C. HQ); three workers, Sa i Khao, Pattani Prov., appressed setae sparsely. S. Thailand, 25.viii.1998, (Sk. Yamane); six workers Body usually bicolored with head, mesosoma, petiole, and two intermediate workers, PSU forest, Hat Yai, postpetiole and ﬁrst gater yellow and with remains of Songkhla Prov., 22.x.2011 (dead twig) (TH11-SKY- gaster brown to black. 168) (Sk. Yamane): three workers, Khlong Saeng, Ratchaprapha, Surat Thani, 15.x.2011 (dead twig on Comments: In the worker this species can be distin- tree) (TH11-SKY-139) (Sk. Yamane). guished from all other members of the C. quadriruga group by the distinct compound eyes, propodeal spira- CREMATOGASTER RETICULATA HOSOISHI, 2009 cles small and apart from metapleural gland bulla, and petiole with subparallel sides in dorsal view. This species (FIGS 116, 117) is similar to C. suehiro, but can be distinguished from Crematogaster (Orthocrema) reticulata Hosoishi, 2009: it by longitudinal rugulae on clypeus not extending to 261; Holotype worker, Ulu Gombak, Selangor, Malay- posterior clypeal margin, bicolored body, petiole with sia, 27.xi.2005 (SH05-Mal-01) (S. Hosoishi) (KUEC, ex- subparallel sides in dorsal view. amined) and ten paratype workers, same data as Specimens from northern part of Thailand (Chiang holotype (BMNH, MCSN, MCZC, MHNG, NHMB, Mai) differ from other specimens in having the devel- examined). oped rugulae on promesonotum, slightly large propodeal Crematogaster reticulata; Blaimer, 2012c: 55 [Com- spiracles and slightly broader petiole with sculptured bination in C.(Orthocrema)]. surface. Single specimen from Laos has erect longer setae on pronotal shoulders medially. Some speci- Worker measurements (n = 7): HW 0.40–0.46; HL 0.40– mens from Bokor National Park, Cambodia differ in 0.46; CI 96–103; SL 0.36–0.42; SI 84–91; EL 0.10– having yellow-colored gaster. 0.11; PW 0.27–0.30; WL 0.46–0.57; PSL 0.09–0.14; PtL 0.15–0.17; PtW 0.12–0.14; PtH 0.11–0.13; PpL 0.09– Distribution and biology: This species is known from 0.12; PpW 0.14–0.15; PtHI 67–76; PtWI 75–87; PpWI Laos, Cambodia, Thailand and Indonesia (N. Sumatra) 122–155; WI 107–111. (Fig. 52). This species inhabits disturbed to developed forests, and nests in dead twigs on trees. General description of worker: Workers monomorphic, but intermediate workers as large as queen (see below). Material examined: CAMBODIA: six workers, Head subquadratic in full-face view. Mandibles with Permanental Sample Plots, Kampong Thom, 9.i.2010 four teeth arranged at an equal distance, apical and (SH10-Cam-20) (S. Hosoishi); two workers, Commu- subapical teeth large, basal two teeth smaller. Ante- nity forest, Kampong Chhnang, 27.xi.2010 (bamboo) rior clypeal margin weakly convex or almost straight (SH10-Cam-198) (S. Hosoishi); one worker, Kampong in medial portion. Compound eyes distinctly project- Thom, 20.xi.2010 (TUS arboreal) (S. Hosoishi); 12 ing beyond lateral margins of head in full-face view. workers, Bokor National Park (870 m alt.), Kampot, Scapes reaching posterolateral corners of head. 9–11.xii.2011 (Winkler 50 cm × 50 cm quadrat) (S. Pronotal collar with almost straight anterior margin Hosoishi & S.-H. Park); INDONESIA: six workers, Pulau in dorsal view, distinctly lower than pronotum in lateral Weh, off Banda Aceh, Aceh, Sumara (disturbed forest), view. Pronotal dorsum with ridges laterally. Mesonotal 27.xi.2012 (dead leaves on tree) (SU12-SKY-208) (Sk. dorsum with lateral ridges. Pronotum and mesonotum Yamane); LAOS: one worker, Phang Dang Vil., Pak- in lateral view not clearly forming continuous dorsal Gnam Distr., Vientinane Prov. (350 m alt.), 13.vi.2010 outline. Metanotal groove in dorsal view transverse, (leaf litter) (Sk. Yamane); THAILAND: two workers, almost straight in median portion, forming deep con- Campus of Chiang Mai Univ., Chiang Mai Prov., N. cavity that is laterally margined by lamellate ridges. Thailand, 10.vi.2001, (Eg01-TH-165) (K. Eguchi); one Propodeal spiracles elliptical, situated at posterolateral worker, Doi Chiang Dao, Chiang Mai Prov., N. Thai- corners of propodeum, apart from metapleural gland land, 9.vi.2001, (Eg01-TH-140) (K. Eguchi); one worker, bullae. Propodeal spines developed, longer than diam- Sakaerat lowland forest (DEF), Nakornratchasima, NE eter of propodeal spiracles, in dorsal view directed Thailand, 10.vii.1999, (Sk. Yamane); 15 workers, posteriad. Residental area, Bang Khean Distr., Bangkok, C. Thai- Petiole in dorsal view with parallel sides and angulate land, 25.viii.2003, (TH03-SKY-102) (dry dead wood) (Sk. shoulders anteriorly, longer than wide. Posterior portion Yamane); three workers, Lum Jang Wat, E. Thai., of petiole with short process that is slightly higher than 7.xii.2005, (SH05-Tha-51) (S. Hosoishi); three workers, posterior margin of petiole disc in lateral view. Maegar, Phayao, 30.iii.1990, (no collector’s name); four Subpetiolar process developed as acute process. workers, Kaeng Krachan NP (320 m alt.), Phetchaburi Postpetiole in lateral view with weakly convex dorsum, Prov. (ex suspended dead twig) (TH14-SKY-10) (Ban as high as petiole, in dorsal view slightly wider than

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 596 S. HOSOISHI AND K. OGATA petiole, weakly bilobed posteriorly but without longi- of stout long setae posteriorly. Postpetiole with four tudinal sulcus. Subpostpetiolar process developed as pairs of stout long setae on disc anterodorsally, acute process. anterolaterally and posteriorly. Integument essentially sculptured. Dorsal surface of head sculptured reticulately. Mandibles with Comments: In the worker this species can be distin- feeble rugulae and smooth interspaces. Clypeus guished from all other members of the C. biroi group weakly sculptured with longitudinal rugulae; longer by the distinct compound eyes, strongly sculptured dorsal rugulae extending to posterior clypeal margin. surface of head, and petiole squared without angulate Anterolateral shoulders of pronotum with rugulae. anterolateral corners. This species is very similar to Lateral surface of pronotum sculptured in higher portion, C. schimmeri, but can be distinguished from it by the but relatively smooth and shining in lower portion. propodeal spiracles apart from metapleural gland bulla, Pronotum and mesonotum with longitudinal rugulae petiole squared without angulate anterolateral corners. and sculptured interspaces. Mesopleura sculptured, but relatively smooth in central areas. Rugula on higher Distribution and biology: This species is known from portion of mesopleura extending to small pit of S. Thailand and Malaysia (Peninsula and Borneo) and mesothoracic spiracles. Dorsal surface of propodeum Brunei (Fig. 51). This species inhabits developed forests, sculptured. Dorsal and lateral surfaces of petiole sculp- and nests in dead twigs or in dead leaves on trees (SH tured. Dorsal and lateral surfaces of postpetiole collections from Ulu Gombak). sculptured. Standing pilosity sparse. Dorsal face of head with Material examined: BRUNEI: one worker, Tasek several pairs (c. 9) of erect and stout long setae, and Merimbun, 12.ii.1999. (Eg99-BOR-032) (K. Eguchi); MA- short and decumbent setae sparsely. Clypeus with two LAYSIA: two workers, Lambir Hills N. P. Miri, Sarawak, pairs of long setae in anterior portion, one directed 20.x.2005. (CH000512) (C. Handa); six workers, Ulu upward and the other downward. Anterior clypeal Gombak, Selangor, 7.iii.2009 (SH09-Mal-37) (S. margin with one pair of long setae medially and some Hosoishi); THAILAND: six workers, Evergreen For., pairs (three to four) of short setae laterally. Scapes with Khlong Naka WS., Ranong Prov., S. Thailand, decumbent to appressed setae. Mesosoma with five pairs 12.viii.2009 (WJT09-TH2046) (W. Jaitrong); four workers, of long erect and stout setae (ps1PN, ps2PN, psaMN, Papra Stn., Khao Nan N. P., Nakhon, S. Thammarat pspMN, and ps2PS) that are much longer than other Prov., S. Thailand, 13.iii.2007 (TH07-SKY-24) (W. erect setae. Posterolateral tubercles of petiole poste- Jaitrong); four workers and two intermediate workers, riorly with one pair of stout long setae. Postpetiole with Datfa Waterfall, Tai Rom Yen National Park, Surat three pairs of stout long setae on disc anterodorsally, Thani Prov., 12.x.2011 (knot of live tree) (TH11-SKY- anterolaterally and posteriorly. Fourth abdominal tergite 062) (Sk. Yamane). with several pairs (c. 12) of erect and stout setae, and short appressed setae sparsely. CREMATOGASTER SCHIMMERI FOREL, 1912a Body yellow-brown. All flagellar segments yellow. (FIG. 118) Intermediate worker measurements (n = 2): HW 0.66– Crematogaster schimmeri Forel, 1912a: 69; syntype 0.75; HL 0.63–0.69; CI 105–109; SL 0.49–0.54; SI 72– workers, Pilam, Taiwan (H. Sauter) (MHNG, exam- 74; EL 0.16–0.18; PW 0.49–0.56; WL 0.80–0.94; PSL ined). One syntype worker (middle specimen of three 0.16–0.18; PtL 0.28–0.32; PtW 0.23–0.26; PtH 0.18– on one pin) in MHNG here designated Lectotype. 0.23; PpL 0.17–0.20; PpW 0.27–0.30; PtHI 64–72; PtWI Crematogaster schimmeri; Emery, 1922: 132 [Com- 81–82; PpWI 150–159; WI 115–117. bination in C.(Decacrema)]. Crematogaster schimmeri; Terayama, 2009: 177 [Com- Description of intermediate worker: With worker char- bination in C.(Decacrema)]. acter conditions, except as follows. Crematogaster schimmeri; Hosoishi & Ogata, 2009a: Head subquadratic. Three ocelli present. 1 [Combination in C.(Orthocrema)]. Mesonotum highly convex in lateral view. Pronotum Crematogaster schimmeri; Blaimer, 2012c: 55 [Com- not clearly forming same dorsal outline with mesonotum bination in C.(Orthocrema)]. in lateral view, but posterior face forming oblique slope Crematogaster schimmeri; Peeters et al., 2013: 258, to metanotal groove. Propodeal spiracles oval. figs 2–4 [External and internal morphology of inter- Subpetiolar process developed as small tubercle. mediate worker]. Mesosoma with nine to 11 pairs of erect and stout long setae; one ps1PN and one to two ps2PN, five to Worker measurements (n = 5): HW 0.44–0.48; HL 0.45– six pairs on mesonotal ridges, two pairs on propodeal 0.49; CI 96–98; SL 0.38–0.40; SI 79–87; EL 0.12– spines. Posterolateral tubercles of petiole with two pairs 0.13; PW 0.29–0.32; WL 0.50–0.54; PSL 0.08–0.10; PtL

0.14–0.17; PtW 0.15–0.17; PtH 0.11–0.14; PpL 0.11– pairs (three to four) of short setae laterally. Scapes with 0.12; PpW 0.16–0.18; PtHI 75–82; PtWI 88–107; PpWI decumbent to appressed setae. Mesosoma with ﬁve pairs 142–150; WI 107–113. of long erect and stout setae (ps1PN, ps2PN, psaMN, pspMN, and ps2PS) that are much longer than other General description of worker: Workers monomorphic. erect setae. Posterolateral tubercles of petiole poste- Head subquadratic in full-face view. Mandibles with riorly with one pair of stout long setae. Postpetiole with four teeth arranged at an equal distance, apical and three pairs of stout long setae on disc anterodorsally, subapical teeth large, basal two teeth smaller. Ante- anterolaterally and posteriorly. Fourth abdominal tergite rior clypeal margin weakly convex in medial portion. with several pairs (c. 12) of erect and stout setae, and Compound eyes distinctly projecting beyond lateral short appressed setae sparsely. margins of head in full-face view. Scapes reaching Body yellow. All ﬂagellar segments yellow. posterolateral corners of head. Pronotal collar with almost straight anterior margin Comments: In the worker this species can be distin- in dorsal view, distinctly lower than pronotum in lateral guished from all other members of the C. biroi group view. Pronotal dorsum with ridges laterally. Mesonotal by the distinct compound eyes, sculptured dorsal surface dorsum with lateral ridges. Pronotum and mesonotum of head, and petiole squared with angulate anterolateral in lateral view not clearly forming continuous dorsal corners. This species is similar to C. reticulata, but outline. Metanotal groove in dorsal view transverse, can be distinguished from it by the propodeal spira- almost straight in median portion, forming deep con- cles touching metapleural gland bulla and squared cavity that is laterally margined by lamellate ridges. petiole with angulate anterolateral corners. Propodeal spiracles oval, situated at posterolateral Terayama (2009) assigned this species to the sub- corners of propodeum, touching metapleural gland genus Decarema by having 10-segmented antennae. bullae. Propodeal spines developed, longer than diam- However, examinations of the type specimens reveals eter of propodeal spiracles, in dorsal view directed that the species has 11-segmented antennae and belongs posteriad. to the subgenus Orthocrema (Hosoishi & Ogata, 2009a). Petiole in dorsal view with parallel sides and angulate shoulders anteriorly, longer than wide. Posterior portion Distribution: This species is known only from the type of petiole with short process that is slightly higher than locality in Taiwan (Fig. 51). posterior margin of petiole disc in lateral view. Subpetiolar process developed as blunt process. CREMATOGASTER STORKI SP. NOV. Postpetiole in lateral view with weakly convex dorsum, as high as petiole, in dorsal view slightly wider than (FIG. 119) petiole, weakly bilobed posteriorly but without longi- Holotype worker. Dumoga-Bone N. P., Utara, Sulawesi, tudinal sulcus. Subpostpetiolar process developed as INDONESIA, Fog. 5, (400 m alt.), 11.ii.1985, BMNH acute process. Plot C (N. Stork) (BMNH). Integument essentially sculptured. Dorsal surface of Paratypes. Five workers, same data as holotype head weakly sculptured reticulately. Mandibles with (CASC, KUEC, MHNG, MBBJ, THNHM). feeble rugulae and smooth interspaces. Clypeus weakly sculptured with longitudinal rugulae; longer rugulae Worker measurements (n = 6): HW 0.47–0.50; HL 0.48– extending to posterior clypeal margin. Anterolateral 0.52; CI 96–100; SL 0.51–0.54; SI 104–113; EL 0.12– shoulders of pronotum with rugulae. Lateral surface 0.13; PW 0.29–0.32; WL 0.60–0.65; PSL 0.17–0.22; PtL of pronotum sculptured in higher portion, but rela- 0.24–0.27; PtW 0.16–0.19; PtH 0.14–0.16; PpL 0.13– tively smooth and shining in lower portion. Pronotum 0.14; PpW 0.18–0.19; PtHI 59–68; PtWI 67–77; PpWI and mesonotum with longitudinal rugulae and sculp- 136–146; WI 100–113. tured interspaces. Mesopleura sculptured, but relatively smooth in central areas. Rugula on higher portion General description of worker: Workers monomorphic. of mesopleura extending to small pit of mesothoracic Head round in full-face view. Mandibles with four spiracles. Dorsal surface of propodeum sculptured. Dorsal teeth arranged at an equal distance, apical and and lateral surfaces of petiole sculptured. Dorsal and subapical teeth large, basal two teeth smaller. Ante- lateral surfaces of postpetiole sculptured. rior clypeal margin convex in medial portion. Com- Standing pilosity sparse. Dorsal face of head with pound eyes distinctly projecting beyond lateral margins several pairs (c. 9) of erect and stout long setae, and of head in full-face view. Scapes exceeding posterolateral short and decumbent setae sparsely. Clypeus with two corners of head. pairs of long setae in anterior portion, one directed Pronotal collar with weakly concave anterior upward and the other downward. Anterior clypeal margin in dorsal view, slightly lower than pronotum margin with one pair of long setae medially and some in lateral view. Pronotal dorsum without distinct ridges

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 598 S. HOSOISHI AND K. OGATA laterally. Mesonotal dorsum with lateral ridges that directed upward at the tip. This species is similar to irregularly extend posteriad to tips of propodeal spines. C. baduvi and C. brunensis, but can be distinguished Pronotum and mesonotum in lateral view forming evenly from them by upward-curved propodeal spines. arched, continuous dorsal outline. Metanotal groove in dorsal view transverse, almost straight in median Distribution: This species is known only from the type portion, forming shallow concavity that is laterally locality in Indonesia (Sulawesi) (Fig. 49). margined by ridges. Propodeal spiracles oval, situated at posteolateral corners of propodeum, apart Etymology: The species name is dedicated to Dr. N. from mepleural gland bullae. Propodeal spines devel- Stork (BMNH), who collected the type material. oped, longer than diameter of propodeal spiracles, in dorsal view strongly divergent, in lateral view curved Material examined: INDONESIA: three workers, upward. Dumoga-Bone N. P., Utara, Sulawesi, 14.ii.1985 (No Petiole in dorsal view with subparallel sides and collector’s name). narrow short peduncle anteriorly, distinctly longer than wide. Posterior portion of petiole with short process that is slightly higher than posterior margin of petiole CREMATOGASTER SUEHIRO TERAYAMA, 1999 disc in lateral view. Subpetiolar process undeveloped. (FIG. 120) Postpetiole in lateral view with strongly convex dorsum, distinctly higher than petiole, in dorsal view as wide Crematogaster suehiro Terayama, 1999: 726, figs. 1–4; as petiole, globular, not bilobed. Subpostpetiolar process holotype worker and two paratype workers, one paratype undeveloped, but venter of postpetiole convex. queen and one intermediate worker, Ishigaki Island, Integument essentially smooth and shining. Dorsal Japan (MNHA, not examined). surface of head smooth and shining. Mandibles with Crematogaster miroku Terayama, 2013: 12, figs 23– feeble rugulae and smooth interspaces. Clypeus gen- 25; Holotype worker and six paratype workers, Higashi- erally smooth and shining, but with one distinct pair son, Kunigami-gun, Okinawa-jima, Okinawa Pref., of longitudinal shorter rugulae. Dorsal and lateral sur- Japan, 3.xi.2012 (H. Takamine) (ITLJ, examined; images faces of pronotum smooth and shining; anterolateral also examined from Yoshitake et al., 2011). Syn. nov. shoulders of pronotum with rugulae. Mesopleura weakly Crematogaster suehiro; Blaimer, 2012c: 55 [Combi- sculptured, but sometimes smooth except for their mar- nation in C.(Orthocrema)]. ginal areas in some specimens (on anterior and central areas). Dorsal surface of propodeum weakly sculp- Worker measurements (n = 6): HW 0.46–0.52; HL 0.47– tured and with rugulae on anterodorsal areas. Dorsal 0.51; CI 96–102; SL 0.46–0.50; SI 94–100; EL 0.12– surface of petiole smooth and shining. Lateral surface 0.14; PW 0.30–0.33; WL 0.54–0.6; PSL 0.11–0.13; PtL of petiole sculptured. Dorsal and lateral surfaces of 0.18–0.20; PtW 0.16–0.18; PtH 0.14–0.15; PpL 0.12– postpetiole smooth and shining. 0.14; PpW 0.17–0.19; PtHI 74–79; PtWI 85–90; PpWI Standing pilosity sparse. Dorsal face of head with 123–158; WI 106–112. three pairs of long erect setae and short appressed setae sparsely. Clypeus with two pairs of long setae in ante- General description of worker: Workers monomorphic. rior portion, one directed upward and the other down- Head subquadratic in full-face view. Mandibles with ward. Anterior clypeal margin with one single long setae four teeth arranged at an equal distance, apical and medially and one pair of long setae laterally. Scapes subapical teeth large, basal two teeth smaller. Ante- with appressed setae. Mesosoma with two pairs of long rior clypeal margin convex in medial portion. Com- erect and stout setae (ps1PN, and psaMN) that are pound eyes distinctly projecting beyond lateral margins much longer than other erect setae and shorter pspMN. of head in full-face view. Scapes reaching posterolateral Posterolateral tubercles of petiole posteriorly with one corners of head. pair of stout long setae. Postpetiole with one pair of Pronotal collar with weakly concave anterior stout long setae on disc posteriorly. Fourth abdomi- margin in dorsal view, distinctly lower than pronotum nal tergite with erect setae sparsely (c. 8), but no de- in lateral view. Pronotal dorsum without distinct cumbent to appressed setae. ridges laterally. Mesonotal dorsum with lateral Body red-brown. Apical two flagellar segments yellow, ridges. Pronotum and mesonotum in lateral view not contracting with other flagellar segments that are brown. clearly forming continuous dorsal outline. Metanotal groove in dorsal view transverse, almost straight in Comments: In the worker this species can be distin- median portion, forming deep concavity that is later- guished from all other members of the C. baduvi group ally margined by lamellate ridges. Propodeal spira- by the smooth and shining surface of mesosoma, cles oval, situated at posterolateral corners of propodeal dorsum with rugulae, and propodeal spines propodeum, touching metapleural gland bullae.

Figures 120–123. Body in lateral view. 120, Crematogaster suehiro, worker [Mt. Omoto, Ishigaki Island, Japan]; 121, Crematogaster sundalandensis, worker [Maliau Basin (riparian forest), Sabah, Borneo, E. Malaysia]; 122, Crematogaster vieti, worker [near Forestry Station, Khe Kem, Pu Mat National Park, Nghe An, Vietnam]; 123, Crematogaster vieti, intermediate worker [near Forestry Station, Khe Kem, Pu Mat National Park, Nghe An, Vietnam].

Propodeal spines developed, longer than diameter Standing pilosity sparse. Dorsal face of head with of propodeal spiracles, in dorsal view directed erect setae sparsely. Clypeus with two pairs of long posteriad. setae in anterior portion, one directed upward and Petiole in dorsal view with subparallel sides and the other downward. Anterior clypeal margin with narrow anteriorly, longer than wide. Posterior portion one pair of long setae medially and short setae later- of petiole without distinct process in lateral view. ally. Scapes with suberect setae. Mesosoma with five Subpetiolar process weakly developed as small process. pairs of long erect and stout setae [ps1PN, ps2PN, Postpetiole in lateral view with weakly convex dorsum, psaMN, pspMN (also one pair of short setae medi- as high as petiole, in dorsal view slightly wider than ally), and ps1PS] that are much longer than other petiole, weakly bilobed posteriorly but without longi- erect setae. Posterolateral tubercles of petiole poste- tudinal sulcus. Subpostpetiolar process undeveloped, riorly with two pairs of stout long setae. Postpetiole but venter of postpetiole convex. with four pairs of setae on disc anterodorsally, Integument essentially smooth and shining. Dorsal anterodorsally and posteriorly. Fourth abdominal tergite surface of head smooth and shining. Mandibles with with erect setae abundantly, and short decumbent setae feeble rugulae and smooth interspaces. Clypeus gen- sparsely. erally smooth and shining, but with one distinct pair Body yellow. All flagellar segments yellow. of longitudinal longer rugulae and one pair of shorter rugulae laterally; longer rugulae extending to posteri- Comments: We have not examined the types of C. or clypeal margin. Dorsal and lateral surfaces of suehiro, but the original description and figures closely pronotum smooth and shining; anterolateral shoul- match the specimens examined. In the worker this ders of pronotum without rugulae. Mesopleura smooth species can be distinguished from all other members and shining. Dorsal surface of propodeum generally of the C. quadriruga group by the distinct compound smooth and shining, but with one longitudinal rugulae eyes, propodeal spiracles small and apart from anteriorly. Dorsal and lateral surfaces of petiole smooth metapleural gland bulla, and petiole tapering anteri- and shining. Dorsal and lateral surfaces of postpetiole orly in dorsal view. This species is similar to C. smooth and shining. quadriruga, but can be distinguished from it by the

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 600 S. HOSOISHI AND K. OGATA longitudinal rugulae on the clypeus extending to the view. Pronotal dorsum without distinct ridges later- bottoms between frontal carinae and petiole tapering ally. Mesonotal dorsum with lateral ridges that ir- anteriorly. regularly extend posteriad to tips of propodeal spines. In the original description of C. miroku, Terayama Pronotum and mesonotum in lateral view forming evenly (2013) separated the species from C. osakensis by the arched, continuous dorsal outline. Metanotal groove in carinate dorsolateral corners of mesonotum and smooth dorsal view transverse, almost straight in median and shining mesopleuron. However, the original de- portion, forming deep concavity that is laterally mar- scription and character states mentioned by him match gined by lamellate ridges. Propodeal spiracles ellipti- well with C. suehiro. Additionally longer scape (SI 96) cal, situated at posterolateral corners of propodeum, in holotype of C. miroku also suggests the affinity with touching to metapleural gland bullae. Propodeal spines C. suehiro (SI 94–100) rather than C. osakensis (SI developed, longer than diameter of propodeal spira- 77–84). In a key to Japanese Crematogaster species cles, in dorsal view directed posteriad. (Terayama, Kubota & Eguchi, 2014), C. miroku was Petiole in dorsal view with subparallel sides and separated from C. suehiro by the petiole with convex narrow anteriorly, longer than wide. Posterior portion sides and slightly broader posteriorly, but he men- of petiole without distinct process in lateral view. tioned ‘Petiole.., widest at posterior end’ in the origi- Subpetiolar process undeveloped. Postpetiole in lateral nal description (Terayama, 2013). Those slight differences view with weakly convex dorsum, as high as petiole, are treated as variation within one species, C. suehiro in dorsal view as wide as petiole, weakly bilobed pos- until additional characters including molecular data teriorly but without longitudinal sulcus. Subpostpetiolar are available. process undeveloped, but venter of postpetiole convex. Integument essentially smooth and shining. Dorsal Distribution and biology: This species is known from surface of head smooth and shining. Mandibles with Southern parts of Japan (Ishigaki Island, Okinawa feeble rugulae and smooth interspaces. Clypeus gen- Island) (Fig. 52). This species inhabits developed forests, erally smooth and shining, but with one pair of lon- and nests in dead twigs on trees. gitudinal rugulae; rugulae not extending to posterior clypeal margin. Dorsal and lateral surfaces of pronotum Material examined: JAPAN: six workers, Mt. Omoto, smooth and shining; anterolateral shoulders of pronotum Ishigaki Island, 10.x.2008 (M. Maruyama & T. without rugulae. Mesopleura smooth and shining. Dorsal Komatsu). surface of propodeum generally smooth and shining, but one pair of rugulae running from metanotal groove to tips of propodeal spines. Dorsal surface of petiole CREMATOGASTER SUNDALANDENSIS SP. NOV. smooth and shining. Lateral surface of petiole gener- (FIG. 121) ally smooth, without longitudinal rugulae. Dorsal and Holotype worker. Parapat (900 m alt.), Danau Toba, N. lateral surfaces of postpetiole smooth and shining. Sumatra, INDONESIA, 19.viii.2002 (SU02-SKY-73) (Sk. Standing pilosity sparse. Dorsal face of head with Yamane) (MBBJ). suberect setae sparsely. Clypeus with two pairs of long Paratypes. Seven workers, same data as hotlotype setae in anterior portion, one directed upward and the (BMNH, CASC, KUEC, MHNG, SKYC, THNHM). other downward. Anterior clypeal margin with one pair of long setae medially and short setae laterally. Scapes Worker measurements (n = 8): HW 0.44–0.53; HL 0.47– with suberect setae mixed with two to three long setae. 0.55; CI 92–100; SL 0.42–0.49; SI 92–96; EL 0.12– Mesosoma with four pairs of long erect and stout setae 0.14; PW 0.29–0.35; WL 0.56–0.64; PSL 0.08–0.11; PtL (ps1PN, psaMN, pspMN, and ps1PS) that are much 0.17–0.21; PtW 0.15–0.16; PtH 0.12–0.15; PpL 0.11– longer than other erect setae. Posterolateral tuber- 0.13; PpW 0.16–0.19; PtHI 70–78; PtWI 80–88; PpWI cles of petiole posteriorly with two pairs of stout setae. 133–146; WI 106–107. Postpetiole with three pairs of setae on disc anteriorly, posteriorly, and laterally. Fourth abdominal tergite General description of worker: Workers monomorphic. with erect setae sparsely, and short decumbent setae Head subquadratic in full-face view. Mandibles with sparsely. four teeth arranged at an equal distance, apical and Body bicolored, head and gaster with brown, subapical teeth large, basal two teeth smaller. Ante- mesosoma, petiole and postpetiole with yellow. All ﬂa- rior clypeal margin convex in medial portion. Com- gellar segments yellow. pound eyes distinctly projecting beyond lateral margins of head in full-face view. Scapes reaching posterolateral Comments: In the worker this species can be distin- corners of head. guished from all other members of the C. quadriruga Pronotal collar with weakly concave anterior margin group by the distinct compound eyes, V-shaped in dorsal view, distinctly lower than pronotum in lateral metanotal groove in lateral view, large propodeal

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 SYSTEMATICS AND BIOGEOGRAPHY OF ASIAN 601 spiracles touching metapleural gland bulla, long General description of woker: Workers weakly poly- propodeal spines (PSL 0.08–0.11), and bicolored body. morphic in size. This species is similar to C. philippinensis, but can Head subquadratic in full-face view. Mandibles with be distinguished from it by the scape with two to three four teeth arranged at an equal distance, apical and erect setae and decumbent setae, and bicolored body. subapical teeth large, basal two teeth smaller. Ante- rior clypeal margin weakly convex in medial portion. Distribution: This species is known from Malaysia Compound eyes distinctly projecting beyond lateral (Borneo) and Indonesia (Sumatra) (Fig. 52). margins of head in full-face view. Scapes reaching posterolateral corners of head. Etymology: The speciﬁc name refers to the regions which Pronotal collar with almost straight anterior margin the material is collected. in dorsal view, distinctly lower than pronotum in lateral view. Pronotal dorsum with distinct ridges laterally. Material examined: INDONESIA: one worker, Ulu Mesonotal dorsum with lateral ridges that irregular- Gadut, nr. Padang, W. Sumatra, 27–30.viii.1985 (Sk. ly extend posteriad to tips of propodeal spines. Pronotum Yamane); MALAYSIA: seven workers, Maliau Basin (ri- and mesonotum in lateral view not clearly forming con- parian forest), Sabah, Borneo (300 m alt.), 10.xi.2011 tinuous dorsal outline. Metanotal groove in dorsal view (ex carton cover) (SB11-SKY-38) (Sk. Yamane). transverse, almost straight in median portion, forming deep concavity that is laterally margined by lamellate ridges. Propodeal spiracles oval, situated at CREMATOGASTER UDO FOREL, 1905 posterolateral corners of propodeum, apart from or Crematogaster sordidula var. udo Forel 1905: 20; worker, touching metapleural gland bullae. Propodeal spines Tjompea, Java, Indonesia [types not found in MHNG]. developed, longer than diameter of propodeal spira- Crematogaster udo; Emery, 1922: 132 [Raised to cles, in dorsal view directed posteriad. species]. Petiole in dorsal view with parallel sides and angulate Crematogaster udo; Emery, 1922: 132 [Combina- shoulders anteriorly, longer than wide. Posterior portion tion in C.(Orthocrema)]. of petiole with short process that is slightly higher than Crematogaster udo; Blaimer, 2012c: 55 [Combina- posterior margin of petiole disc in lateral view. tion in C.(Orthocrema)]. Subpetiolar process developed as acute process. Postpetiole in lateral view with weakly convex dorsum, Comments: We have not been able to examine the type as high as petiole, in dorsal view as wide as petiole, material of C. udo. Forel (1905) referred the affinity globular, not bilobed. Subpostpetiolar process devel- with C. fritzi Emery in the original description, whereas oped as acute process. Menozzi’s key (1935) clearly separated C. fritzi from Integument essentially smooth and shining. Dorsal other Orthocrema species, including C. udo, by having surface of head generally smooth and shining, but two pairs of hairs on mesosomal dorsum. Taxonomic weakly with rugulae on surrounding region of antennal relationship of C. udo with C. fritzi will remain un- sockets. Mandibles with feeble rugulae and smooth resolved until the type material is examined. interspaces. Clypeus generally smooth and shining, but with two distinct pairs of longitudinal rugulae; rugulae not extending to posterior clypeal margin. Anterolateral CREMATOGASTER VIETI SP. NOV. shoulders of pronotum with rugulae. Dorsal surface of (FIGS 122, 123) pronotum with longitudinal rugulae. Lateral surface Holotype worker. Y Linh Ho, Sa Pa, Lao Cai, VIETNAM, of pronotum smooth and shining. Mesopleura weakly Small fragment of forest (c. 1100 m alt.), 1.v.2002 (Eg02- sculptured, but relatively smooth on central areas. VN-225) (K. Eguchi) (IEBR). Rugula on higher portion of mesopleura extending to Paratypes. Two workers, same data as holotype small pit of mesothoracic spiracles. Dorsal surface of (ACEG, KUEC); three workers, Y Linh Ho, Sa Pa, Lao propodeum with rugulae. Dorsal surface of petiole Cai, VIETNAM, Small fragment of forest (c. 1100 m smooth and shining. Lateral surface of petiole weakly alt.), 1.v.2002 (Eg02-VN-215) (K. Eguchi) (BMNH, CASC, sculptured. Dorsal and lateral surfaces of postpetiole THNHM). smooth and shining. Standing pilosity sparse. Dorsal face of head with Worker measurements (n = 9): HW 0.42–0.70; HL 0.45– two to three pairs of erect and stout long setae, and 0.64; CI 92–113; SL 0.37–0.51; SI 72–88; EL 0.10– short and decumbent setae sparsely. Clypeus with two 0.16; PW 0.26–0.39; WL 0.54–0.75; PSL 0.07–0.14; PtL pairs of long setae in anterior portion, one directed 0.17–0.24; PtW 0.13–0.20; PtH 0.12–0.18; PpL 0.11– upward and the other downward. Anterior clypeal 0.16; PpW 0.15–0.21; PtHI 74–86; PtWI 77–88; PpWI margin with one pair of long setae medially and some 120–154; WI 95–111. pairs of short setae laterally. Scapes with suberect to

© 2016 The Linnean Society of London, Zoological Journal of the Linnean Society, 2016, 176, 547–606 602 S. HOSOISHI AND K. OGATA decumbent setae. Mesosoma with four pairs of long erect Distribution: This species is known from Vietnam and stout setae (ps1PN, psaMN, pspMN, and ps1PS) (Fig. 51). that are much longer than other erect setae. Posterolateral tubercles of petiole posteriorly with one Etymology: The species name is dedicated to Dr. Bui pair of stout long setae. Postpetiole with four pairs of T. Viet, who helped with field surveys in Vietnam. stout long setae on disc anterodorsally, anterolaterally and posteriorly. Fourth abdominal tergite with erect Material examined: VIETNAM: one worker, Tam Dao and stout setae sparsely, and short decumbent setae 1000 m alt., Vinh Phuc Prov., N. Vietnam, 8.viii.1998 sparsely. (Sk. Yamane); one worker, Tam Dao 1000 m alt., Vinh Body yellow. All flagellar segments yellow. Phuc Prov., N. Vietnam, 9.viii.1998 (Sk. Yamane); two workers, Tam Dao 1000 m alt., Tam Duong Dist., Vinh Intermediate worker measurements (n = 1): HW Prov., 6.xi.2001 (K. Ogata); four workers, Ba Vi 460 0.68; HL 0.70; CI 97; SL 0.54; SI 79; EL 0.19; PW m alt., Ba Vi Dist., Ha Tay Prov., 11.xi.2001 (K. Ogata); 0.47; WL 0.95; PSL 0.19; PtL 0.32; PtW 0.25; PtH 0.24; two workers, Ba Vi 670 m alt., Ba Vi Dist., Ha Tay PpL 0.20; PpW 0.28; PtHI 75; PtWI 78; PpWI 140; WI Prov., 12.xi.2001 (K. Ogata); four workers, Tu Lung, 112. Mai Chau Dist., Hoa Binh Prov., 27.xi.1999 (K. Ogata); three workers, Small fragment of forest (c. 1100 m alt.), Y Linh Ho, Sa Pa, Lao Cai, 1.v.2002 (Eg02-VN-215) Description of intermediate worker: With worker char- (K. Eguchi); three workers, Small fragment of forest acter conditions, except as follows. (c. 1100 m alt.), Y Linh Ho, Sa Pa, Lao Cai, 1.v.2002 Vestigial lateral ocelli present. (Eg02-VN-225) (K. Eguchi); four workers, M. Nghe An, Mesonotum highly convex in lateral view. Mesonotal Pu Hoat, Lung Khung (640 m alt.), 5.xi.1999 (code- dorsum without lateral ridges. Pronotum not clearly 011) (T. V. Bui); one worker, Chua Yen Tu (720– forming same dorsal outline with mesonotum in lateral 845 m alt.), Quang Ninh, 19.v.2004 (K. Eguchi); two view, but posterior face forming oblique slope to workers and one intermediate worker, near Forestry metanotal groove. Propodeal spiracles touching Station (253 m alt.), Khe Kem, Pu Mat N. P., Nghe metapleural gland bulla. An, 15.iii.2006 (Eg15iii06-03) (K. Eguchi). Petiole wth subparallel sides in dorsal view, tapering anteriorly. Subpetiolar process developed as blunt Species excluded from the subgenus Orthocrema in Asia. process. Dorsal surface of pronotum smooth and shining. CREMATOGASTER CRASSICORNIS EMERY, 1893 Dorsal surface of mesonotum with rugulae anteriorly Crematogaster crassicornis Emery, 1893: 265, pl. 2, figs and laterally. Mesopleura generally smooth, but oblique 4, 5; syntype workers, Manila, Philippines (M. E. Simon) sulcus running from anterior areas. (MCSN, MHNG, examined). Anterodorsal surface of propodeum with rugulae Crematogaster crassicornis; Emery, 1922: 132 [Com- reticulately. bination in C.(Orthocrema)]. Anterior clypeal margin with one single and one pair Crematogaster crassicornis; Blaimer, 2012c: 55 [Com- of long setae on median portion and some pairs of short bination in C.(Orthocrema)]. setae laterally. Mesosoma with several pairs of long setae on promesonotum, one ps1PS. Posterolateral tubercles of petiole with two to three pairs of stout long Comments: Examination of syntype workers in MCSN setae posteriorly. Fourth abdominal tergite with suberect and MHNG reveals that C. crassicornis does not belong to decumbent setae abundantly. to the subgenus Orthocrema. Although this species has two-segmented antennal club, characteristic features include: anterolateral margins of clypeus protruded an- Comments: In the worker this species can be distin- teriorly; petiole without node-like process posteriorly guished from all other members of the C. biroi group in lateral view; postpetiole bilobed with longitudinal by the distinct compound eyes, generally smooth dorsal median sulcus. These features are characteristic of the surface of head, petiole with subparallel sides in dorsal subgenus Crematogaster and we consider C. crassicornis view, and erect setae on body tapering distally. This to be referred to that subgenus. species is very similar to C. osakensis, but distinguished from it by the thick propodeal spines and petiole with subparallel sides. CREMATOGASTER PAULI EMERY, 1901 This species corresponds to sp. 36 of SKY (Eguchi Crematogaster pauli Emery, 1901: 575, fig. A; syntype et al., 2005) and sp. eg-1, sp. eg-4 by Eguchi et al. workers, Salabanka, Sulawesi, Indonesia (MCSN, (2011). MHNG, examined).

Crematogaster pauli; Emery, 1922: 132 [Combina- Blaimer BB. 2010. Taxonomy and natural history of the tion in C.(Orthocrema)]. Crematogaster (Decacrema) group (Hymenoptera: Formicidae) Crematogaster pauli; Blaimer, 2012c: 55 [Combina- in Madagascar. Zootaxa 2714: 1–39. tion in C.(Orthocrema)]. Blaimer BB. 2012a. Untangling complex morphological variation: taxonomic revision of the subgenus Crematogaster Comments: Examination of syntype workers in MCSN (Oxygyne) in Madagascar, with insight into the evolution and and MHNG reveals that C. pauli does not belong to biogeography of this enigmatic ant clade (Hymenoptera: the subgenus Orthocrema. Although this species has Formicidae). Systematic Entomology 37: 240–260. two-segmented antennal club, characteristic features Blaimer BB. 2012b. Taxonomy and species-groups of the subgenus Crematogaster (Orthocrema) in the Malagasy region include: anterolateral margins of clypeus protruded an- (Hymenoptera, Formicidae). Zookeys 199: 23–70. teriorly; petiole without node-like process posteriorly Blaimer BB. 2012c. A subgeneric revision of Crematogaster in lateral view. These features are characteristic of the and discussion of regional species groups (Hymenoptera: subgenus Crematogaster and we consider C. crassicornis Formicidae). Zootaxa 3482: 47–67. to be referred to that subgenus. Blaimer BB. 2012d. Acrobat ants go global-Origin, evolution and systematics of the genus Crematogaster (Hymenoptera: ACKNOWLEDGEMENTS Formicidae). Molecular Phylogenetics and Evolution 65: 421– 436. We would like to thank Seiki Yamane for collecting Blaimer BB. 2013. Taxonomy of the Crematogaster degeneri- specimens and his continuing support. We also thank species-assemblage in the Malagasy region (Hymenoptera: to the following persons for loans of material or access Formicidae). European Journal of Taxonomy 51: 1–64. to museum collections: Suzanne Ryder and Gabin Broad Bolton B. 1988. Secostruma, a new subterranean tetramoriine (BMNH); Daniel Burckhardt (NHMB); Sándor Csõsz ant genus (Hymenoptera: Formicidae). Systematic Entomol- (HNHM); Bernhard Merz (MHNG); Maria Tavano and ogy 13: 263–270. Roberto Poggi (MCSN); Dominique Zimmermann Bolton B. 1995. A new general catalogue of the ants of the (NHMW). Additional material was received from world. Cambridge, Massachusetts: Harvard University Press. Himender Bharti, Viet Bui, Katsuyuki Eguchi, John 504 pp. Fellowes, Weeyawat Jaitrong, Takashi Komatsu, Bolton B. 2003. Synopsis and classification of Formicidae. Munetoshi Maruyama, Martin Pfeiffer. Thanks are also Memoirs of the American Entomological Institute 71: 1–370. for due to Rosli Hashim (Universiti Malaya, Malay- Bolton B. 2007. Taxonomy of the dolichoderine ant genus sia), Phourin Chhang (Forestry Administration, Cam- Technomyrmex Mayr (Hymenoptera: Formicidae) based on bodia), Wattanachai Tasen (Kasetsart University, the worker caste. Contributions of the American Entomo- Thailand), Venancio Sumarita (National Museum of the logical Institute 35: 1–149. Bolton B. 2014. An online catalog of the ants of the world. Philippines) who provisioned our collecting trips. We Available at http://antcat.org (accessed March 2015). would like to thank the ANeT members for encour- Briggs JC. 1987. Biogeography and plate tectonics. develop- agement. Thanks are also due to Mark Lorenz (Forte ments in palaeontology and stratigraphy, 10. Amsterdam: Inc.) for improving the English. This work was sup- Elsevier. 204 pp. ported in part by JSPS KAKENHI (Grant-in-Aid for Brown WL Jr. 1949. Notes on Chinese ants. 1. Crematogaster Scientific Research (C)) Grant Number 26440221 and Lund. Mushi 20: 37–38. KAKENHI (Grand-in-Aid for Scientific Research (B)) Buren WF. 1958. A review of the species of Crematogaster, Grant Number 26304014. sensu stricto, in North America (Hymenoptera: Formicidae). Part I. 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