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Additional report of Tenebrionidae collected from fruiting bodies of polypores at eight sites, including seven national parks, in Sarawak, with descriptions of nine new species, Basides nakashizukai sp. nov., Basides ornatimarginatus sp. nov., Basides rhinoceros sp. nov., Boletoxenus persimilis sp. nov., Bolitonaeus grimmi sp. nov., Menimus (Menimus) pygmaeus sp. nov., Menimus (Menimus) sphaericus sp. nov., Neomida sarawakensis sp. nov., Pentaphyllus lambirensis sp. nov.; Three synonyms are proposed: Basides flavofasciatus Pic, 1916 = Basides bifasciatus Motschulsky, 1873; Ischnodactylus sexguttatus Gebien, 1925c = Basides trimaculatus Pic, 1916; Platydema sexpictum Kaszab, 1939 = Basides trimaculatus Pic, 1916.

Crematogaster modiglianii Crematogaster modiglianii has a widespread distribution in South-East Asia, but does not occur in Java, while another widespread species, C. seaeardi, occurs also in Java. What could explain the absence of C. modiglianii on Java? Menzel & Blüthgen (2010) revealed parabiotic associations between C. modiglianii and Camponotus (Myrmotarsus) rufifemur Emery, 1900. Camponotus rufifemur is known from the Indochinese Peninsula, Malay Peninsula, Borneo, Sumatra and Philippines, but not distributed in Java (Antmaps, 2021; AntWiki, 2021). The close relationship might have affected the absence of C. modiglianii in Java. Alternately, Java has a different climate from the other islands on the Sunda Shelf (Wilting et al., 2012). The climate of Java is drier than in other parts of the Sunda Shelf (Heaney, 1991). Veron et al. (2007) suggested that open and drier habitat in Java at the time might have affected the species distribution of the mongoose Herpestes species. Land bridges connecting the Sundaic regions might have allowed for the migration of C. modiglianii, but the species did not successfully colonize, probably due to ecological competition with related taxa., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Menzel F, Bluthgen N. 2010. Parabiotic associations between tropical ants: equal partnership or parasitic exploitation? Journal of Animal Ecology 79: 71 - 81.","Antmaps. 2021. Antmaps. Available at: http: // www. antmaps. org / (accessed on 1 March 2021).","Wilting A, Sollmann R, Meijaard E, Helgen KM, Fickel J. 2012. Mentawai's endemic, relictual fauna: is it evidence for Pleistocene extinctions on Sumatra? Journal of Biogeography 39: 1608 - 1620.","Heaney LR. 1991. A synopsis of climatic and vegetational change in South-East-Asia. Climatic Change 19: 53 - 61.","Veron G, Patou M-L, Pothet G, Simberloff D, Jennings AP. 2007. Systematic status and biogeography of the Javan and small Indian mongooses (Herpestidae, Carnovora). Zoologica Scripta 36: 1 - 10."]}

Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus, Pham, Thai Hong (2023): Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia. Zoological Journal of the Linnean Society 198 (3): 901-922, DOI: 10.1093/zoolinnean/zlad005, URL: https://academic.oup.com/zoolinnean/article/198/3/901/7197147

The Crematogaster difformis-subgroup The relationships of the sister-species in the C. difformis -subgroup were variable with respect to their geographical distribution patterns. Crematogaster ampullaris and C. seaeardi display an allopatric distribution, whereas C. difformis and C. tanakai display a sympatric distribution. The closely related C. ampullaris and C. seaeardi illustrate an interesting distribution pattern with C. ampullaris in Sulawesi and C. seaeardi widespread elsewhere in South-East Asia. The estimated dating suggests that C. ampullaris and C. seaeardi diverged approximately 4 Mya (Fig. 4; Table 2). The divergence implies a recent split of these populations and landmass distributions in that geological time (Hall, 2002) suggesting that the ancestor of both species dispersed from the Sundaic region to Sulawesi west-to-east across Wallace’s Line. Sulawesi is considered to have taken its present form due to the collision of different islands c. 15–5 Mya (Lohman et al., 2011). As the last possible connections between Borneo and Sulawesi were during the Pliocene or the Miocene (Morales & Melnick, 1998; Mercer & Roth, 2003), the colonization of Sulawesi by the common ancestor may have occurred much later after the island collision event. The ancestor of C. seaeardi and C. ampullaris may have expanded its range to western Sulawesi (eastern Sunda Shelf) during low sea-level periods and evolved into C. ampullaris. As the Makassar Strait is a deep trench, Borneo and Sulawesi were not connected when sea levels dropped. Their diversification resulted from ‘an inter-island dispersal combined with climatic vicariance event’ (Condamine et al., 2013). Crematogaster difformis and C. tanakai are sympatric in Borneo, and their habitats are spatially close to each other. Crematogaster difformis has an obligate mutualistic relationship with Lecanopteris and Platycerium ferns in canopy trees of lowland rainforests, and C. tanakai inhabits the same nests with C. difformis (Tanaka et al., 2009, 2012; Tanaka & Itioka, 2011). This restricted distribution demonstrates distinct endemism and is, therefore, of conservation significance. Lecanopteris fern species are found in the canopies of lowland tropical rainforests. The preservation of such primary forests will lead to the conservation of the endemic species. Molecular dating indicates that C. difformis and C. tanakai diverged approximately 1 Mya (Fig. 4; Table 2). The low divergence implies a recent speciation. As in C. aurita, the dispersal event and competition can explain similarly the position of C. mucronata, which is endemic to Sumatra (Fig. 4). Klaus et al. (2013) suggested early divergence of freshwater crabs Parathelphusa and subsequent appearance on Sumatra, since much of Sumatra was still submerged during the Late Miocene. In contrast, the ancestor of C. mucronata already appeared on fragmented Sumatra during the Late Miocene. Freshwater crabs require more land area for their habitats, but ants presumably do not require large areas as Yamane (2013) listed 126 ant species having colonized Krakatau Island after eruption. Faunal and genetic similarities between the Malay Peninsula and Sumatran populations have been documented for several taxa (Gorog et al., 2004; Lohman et al., 2011; Leonard et al., 2015), as the two areas were connected at sea level 20–30 m below the present level (Voris, 2000)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Hall R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences 20: 353 - 431.","Lohman DJ, de Bruyn M, Page T, von Rintelen K, Hall R, Ng PKL, Shin H-T, Carvalho GR, von Rintelen T. 2011. Biogeography of the Indo-Australian Archipelago. Annual ReVieae of Ecology, EVolution, and Systematics 42: 205 - 226.","Morales JC, Melnick DJ. 1998. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), as revealed by high resolution restriction site mapping of mitochondrial ribosomal genes. Journal of Human EVolution 34: 1 - 23.","Mercer JM, Roth VL. 2003. The effects of Cenozoic global change on squirrel phylogeny. Science 299: 1568 - 1572.","Condamine FL, Toussaint EFA, Cotton AM, Genson GS, Sperling FAH, Kergoat GJ. 2013. Fine-scale biogeographical and temporal diversification processes of peacock swallowtails (Papilio subgenus Achillides) in the Indo-Australian Archipelago. Cladistics 29: 88 - 111.","Tanaka HO, Inui Y, Itioka T. 2009. Anti-herbivore effects of an ant species, Crematogaster difformis, inhabiting myrmecophytic epiphytes in the canopy of a tropical lowland rainforest in Borneo. Ecological Research 24: 1393 - 1397.","Tanaka HO, Yamane S, Itioka T. 2012. Effects of a ferndwelling ant species, Crematogaster difformis, on the ant assemblages of emergent trees in a Bornean tropical rainforest. Annals of the Entomological Society of America 105: 592 - 598.","Tanaka HO, Itioka T. 2011. Ants inhabiting myrmecophytic ferns regulate the distribution of lianas on emergent trees in a Bornean tropical rainforest. Biology Letters 7: 706 - 709.","Klaus S, Selvandran S, Goh JW, Wowor D, Brandis D, Koller P, Schubart CD, Streit B, Meier R, Ng PKL, Yeo DCJ. 2013. Out of Borneo: Neogene diversification of Sundaic freshwater crabs (Crustacea: Brachyura: Gecarcinucidae: Parathelphusa). Journal of Biogeography 40: 63 - 74.","Yamane SK. 2013. A review of the ant fauna of the Krakatau Islands, Indonesia. Bulletin of the Kitakyushu Museum of Natural History and Human History 11: 1 - 66.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Leonard JA, Den Tex R-J, Hawkins MTR, Munoz-Fuentes V, Thorington R, Maldonado JE. 2015. Phylogeography of vertebrates on the Sunda Shelf: a multi-species comparison. Journal of Biogeography 42: 871 - 879.","Voris HK. 2000. Maps of Pleistocene sea levels in South-East Asia: shorelines, river systems and time durations. Journal of Biogeography 27: 1153 - 1167."]}

The Crematogaster inflata-subgroup Crematogaster aurita is distributed in Indochina, and does not extend southward beyond the Isthmus of Kra. The phylogenetic position of the species was sister to C. onusta within the C. inflata -subgroup. Biogeographic analyses based on the distribution pattern and phylogenetic position of the species suggests that C. aurita appeared as a result of a dispersal event (Fig. 4). During the Pliocene to Pleistocene, terrestrial Indochinese animals might have had many opportunities to disperse to the Sundaic regions and vice versa, since these land areas were connected in the Sunda Shelf during glacial periods. The absence of C. aurita to the south of the Isthmus of Kra presumably resulted from limited dispersal capability and could also be a consequence of competition between related species. The distribution in the Indochinese Peninsula is possibly related to the preference of C. aurita for open, dry habitats. This preference could be explained by savannah corridors that are thought to have spanned Sundaland during glacial periods (Bird et al., 2005) and drier and seasonal habitats are currently present in the Indochinese Peninsula and Java. However, Crematogaster inflata and C. physothorax are sympatric in the Malay Peninsula. Our molecular dating analyses revealed that C. inflata and C. physothorax diverged approximately 6 Mya (Fig. 4; Table 2). This high divergence implies an old split of these populations. It is unclear whether this divergence resulted from a vicariant event or dispersal of the ancestor of both species. The sympatric distribution of these two species in the peninsula might be due to secondary contact after their speciation (Fig. 4). The estimated divergence suggests that the present geographic overlap of the two species is the result of independent colonization events from different geographic centres of origin. Hosoishi & Ogata (2009: figs 25, 28) represented the distribution records of Crematogaster inflata and C. physothorax based on the original descriptions and literature, but the allocations in the figures seem controversial. A record of C. inflata from Myanmar was based on misidentification. Yet extensive field surveys in the Indochinese Peninsula were not able to find C. physothorax (e.g. Khachonpisitsak et al., 2020), whose type locality is Thagata, Tenasserim, Myanmar and presumably this species is not distributed in the Indochinese Peninsula., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 916, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Hosoishi S, Ogata K. 2009. A taxonomic revision of the Asian endemic subgenus Physocrema of the genus Crematogaster (Hymenoptera: Formicidae). Zootaxa 2062: 15 - 36.","Khachonpisitsak S, Yamane S, Sriwichai P, Jaitrong W. 2020. An updated checklist of the ants of Thailand (Hymenoptera, Formicidae). ZooKeys 998: 1 - 182."]}

The Crematogaster Ʋacca-subgroup The phylogenetic relationship between the C. Ʋaccasubgroup and their ancestral state reconstruction suggests that their common ancestor occurred in the Sundaic region and its founder dispersed to Sumatra (Fig. 4). During glacial periods, the populations (C. yamanei) retracted into glacial forest refugia. When environmental conditions became favourable for expansion, the populations recolonized, but in the Sundaic region they were restricted to higher elevation areas due to competition with the C. inflata -subgroup and the C. difformis -subgroup, which were already present in this area at lower elevations. Crematogaster yamanei is endemic to Sumatra, but closely related to C. Ʋacca, which is distributed in the Malay Peninsula and Borneo. Meijaard’s (2004) mammal studies suggested that Sumatra shares elements of its fauna with both the Bornean/Javan and Malayan regions as a result of secondary contact. Those patterns are found in the younger taxon C. yamanei, but not in the older taxon C. mucronata. The restricted distributions of C. mucronata and C. yamanei could have resulted from another speciation event and extirpation of the relatives, respectively., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on page 917, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Meijaard E. 2004. SolVing mammalian riddles: a reconstruction of the Tertiary and Quaternary distribution of mammals and their palaeoenVironments in island South-East Asia. PhD Thesis, Australian National University, 349."]}

IS CREMATOGASTER SUBCIRCULARIS A RELICT SPECIES? Crematogaster subcircularis was revealed to have an interesting distribution pattern. This species is found in mountainous areas (600–1250 m elevation) of the Sundaic region: the Malay Peninsula (Fraser’s Hill, 1000 m), Borneo (Poring, 600–700 m; Crocker Range, 1100 m) and Java (Cikaniki, 1000–1250 m). This distribution suggests that the ancestor retreated to high-altitude forests during the ice age, whereas the other related species (C. modiglianii) remained in lowaltitude forests (0–600 m elevation). Despite a postulated savannah corridor through central Sundaland (Bird et al., 2005) that might have restricted the west– east dispersal of forest species, gallery forests in the Sunda Shelf could have served as corridors for forest species and facilitated the dispersal of C. subcircularis (Fig. 4). Rising sea-levels in the Late Pleistocene may have isolated the three populations. Molecular dating estimates and restricted highland distribution imply that C. subcircularis is a relict species. Several studies have hypothesized that continuous mountain ranges, which span Myanmar, Thailand, Laos, south China and Vietnam, acted as glacial refugia (Brandon-Jones, 1996; Gorog et al., 2004; Iyengar et al., 2005). Assuming that C. subcircularis is a relict with a wide distribution range in the glacial periods, those populations were fragmented into several geographical ones when the sea level was higher, then closely related or sister-species might have retreated to the high-elevation refugia of mainland South-East Asia (Surridge et al., 1999)., Published as part of Hosoishi, Shingo, Maruyama, Munetoshi, Yamane, Seiki, Jaitrong, Weeyawat, Hashim, Rosli, Syaukani, Syaukani, Sokh, Heng, Itioka, Takao, Meleng, Paulus & Pham, Thai Hong, 2023, Multilocus phylogeny and historical biogeography of the Crematogaster inflata-group (Hymenoptera: Formicidae) in South-East Asia, pp. 901-922 in Zoological Journal of the Linnean Society 198 (3) on pages 917-918, DOI: 10.1093/zoolinnean/zlad005, http://zenodo.org/record/8141964, {"references":["Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular South-East Asia during the Last Glacial Period: a savanna corridor in Sundaland? Quaternary Science ReVieaes 24: 2228 - 2242.","Brandon-Jones D. 1996. The Asian Colobinae (Mammalia, Cercopithecidae) as indicators of Quaternary climatic changes. Biological Journal of the Linnean Society 59: 327 - 350.","Gorog AJ, Sinaga MH, Engstrom MD. 2004. Vicariance or dispersal? Historical biogeography of three Sunda shelf murine rodents (Maxomys surifer, Leopoldamys sabanus and Maxomys aehiteheadi). Biological Journal of the Linnean Society 81: 91 - 109.","Iyengar A, Babu N, Hedges S, Venkataraman A, Maclean N, Morin PA. 2005. Phylogeography, genetic structure and diversity in the dhole (Cuon alpinus). Molecular Ecology 14: 2281 - 2297.","Surridge AK, Timmins RJ, Hewitt GM, Bell DJ. 1999. Striped rabbits in South-East Asia. Nature 400: 726 - 726."]}

2. Colocasiomyia oligochaeta species subgroup, new Diagnosis. Antennal arista apically truncate, stick-shaped (ch.4-1; Fig. 3D, E). Proclinate, orbital setae absent (ch.8- 1; Fig. 2D, E). Anterior, dorsocentral setae absent (ch.26-2; Fig. 6D, E). Patch of supracervical setae present over basal portion of dorsolateral, tentorial apodeme (ch.13-1; Fig. 4D, E). Dorsolateral, tentorial apodemes distally strongly curved outward (ch.14-1; Fig. 4D, E). Palpus without prominent seta (ch.17-2; Fig. 5D 1, E 1). 5x index> 4.0 (ch.32-2; Fig. 8D 2, E 2). Hind tibia apicoventrally with two stout setae (ch.40-1; Fig. 9D 3, E 3). Epandrium with neither pubescence nor setae (ch.53-1, ch.54-1; Fig. 11-2D, E). Shared characters. Antennal pedicel grayish brown (Fig. 14D, E). Eye with fine, sparse, interfacetal setulae (ch.7-0; Fig. 2D 1, E 1). Outer, vertical setae absent (ch.9-1; Fig. 4D, E). Inner (convergent) vertical setae dislocated outside line of ipsilateral orbitals; posterior reclinate orbitals longest on head (Fig. 2D 2, E 2). Upper, postocular setae upright or up-curved (ch.10-1; Fig. 4D, E). Dorsomedial, tentorial apodeme reaching 1/5 to vertex (Fig. 4D, E). Facial carina wider than 1st flagellomere, as long as pedicel + 1st flagellomere (Fig. 2D 2, E 2). Clypeus thicker than 1/2 length of cibarium (Fig. 5D 1, E 1). Cibarial, medial sensilla 2 per side; posterior sensilla absent (ch.21-3; Fig. 5D 2, E 2). Labellum with 7 pseudotracheae (ch.24-0; Fig. 5D 1, E 1). Postpronotal lobe with 1 prominent seta (ch.25- 2; Fig. 6D, E). Prescutellar, acrostichal setae absent (ch.27-1; Fig. 6D, E). Only one prominent katepisternal seta longer than acrostichal setulae (ch.29-2; Fig. 7D, E). Costal setae in middle row from 2nd costal section to proximal portion of 3rd section all weak, trichoid; sparse setae in upper row stout, longer, trichoid (ch.30-2, ch.31-1; Fig. 8D 1, E 1). Patch covered with only minute pubescence present on anterodorsal portion of fore tarsomere I (ch.33-1; Fig. 9D 1, E 1). Fore tarsomere II with 1 large and 1 small pegs; latter 1/3 as long as former (ch.37-1; Fig. 9D 1, E 1). Mid tarsomere I as long as tarsomeres II, III and IV combined; hind I slightly longer than II+III+IV (Fig. 9D 2,3, E 2,3). Claws as long as tarsomere V (ch.43-1; Fig. 9D, E). Abdominal tergites blackish brown (Fig. 14D, E). All sternites longer than wide and I fused to II in both sexes (ch.45-2; Fig. 10D, E); male V not fused to VI, and VI completely bilobed (ch.46-0, ch.47-1; Fig. 10D 1, E 1); female sternite VII more or less convex on distal margin (ch.51-2; Fig. 10D 2, E 2). Female tergite VII mid-dorsally separated into 2 triangular, lateral plates (ch.52-0). Epandrial, ventral lobe differentiated (ch.56-1; Fig. 11-2D, E). Surstylus absent (ch.59-4; Fig. 11-2D, E). Hypandrial phragma anterior, strongly sclerotized rod-like portion (ch.68-2; Fig. 11-2D, E). Membranous aedeagus long, tube-like (ch.70-1; Fig. 11-2D, E). Phallal sheath not pubescent (ch.74-0; Fig. 11-2D, E). Phallapodeme articulated to and not shorter than phallal sheath (ch.75-0, ch.76-0; Fig. 11-2D, E). Hypoproct partly pubescent (ch.80-1; Fig. 12D 1, E 1). Epigynium dorsally separated into 2 lateral lobes (ch.81-1; Fig. 12D 1, E 1). Perineal membrane with patch of dense, distinct warts (ch.84-1; Fig. 12E). Hypogynial valve with trichoid sensilla on only distal portion; distal, narrow portion as long as basal, broad portion (Fig. 12D, E). Remarks. This species subgroup is well demarcated by a number of non-homoplastic synapomorphies (Fig. 13) defined as the diagnostic characters, especially the peculiar chaetotaxy that the proclinate orbital, outer vertical, and anterior dorsocentral setae are lacking. Such an anomaly from the basic chaetotaxy is rare in the Drosophilidae, but is seen in a few species of Colocasiomyia: C. crassipes (de Meijere) and C. micheliae of the crassipes group lack the outer vertical and dorsocentral setae (Grimaldi 1991; Yafuso et al. 2000); and C. ecornuta Toda & Takano of the cristata group lacks the outer vertical setae (Takano et al. 2021). Included taxa. Colocasiomyia oligochaeta sp. nov., and C. grimaldii sp. nov., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on pages 225-226, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["Grimaldi, D. A. (1991) Systematics of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae): cladistics, a new generic synonym, new record, and a new species from Nepal. Entomologica Scandinavica, 22, 417 - 426. https: // doi. org / 10.1163 / 187631291 X 00219","Yafuso, M., Sultana, F., Sasaki, Y. & Toda, M. J. (2000) A new species of Colocasiomyia de Meijere (Diptera, Drosophilidae) from North Sulawesi, Indonesia. Entomological Science, 3, 115 - 119.","Takano, K. T., Gao, J. - J., Hu, Y. - G., Li, N. - N., Yafuso, M., Suwito, A., Repin, R., Pungga, R. A. S., Meleng, P. A., Kaliang, C. H., Chong, L. & Toda, M. J. (2021) Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species. Zootaxa, 5079 (1), 1 - 70. https: // doi. org / 10.11646 / zootaxa. 5079.1.1"]}

5) Colocasiomyia grimaldii Gao & Toda, sp. nov. (Figs 2–12E, 14E) Colocasiomyia sp.4 aff. nepalensis: Sultana et al., 2006: 694. Diagnosis. Antennal arista approximately 1.6 times as long as first flagellomere (Fig. 3E). Distance between antennal scapes narrower than scape diameter (Fig. 2E 2). Acrostichal setulae in 8 rows (Fig. 6E). All legs dark grayish yellow except for mid and hind tarsomeres I–IVs grayish yellow (Fig. 14E). Cercus roundish on ventral margin (Fig. 11- 2E). Long, tube-like, membranous aedeagus ornamented with dense, stronger spinules (Fig. 11-2E). Hypogynial valves with roundish, dorsal flaps not fused to each other at distal end of basal portion (Fig. 12E). Description (♁ and ♀). Head. Antennal, first flagellomere approximately 1.3 times as long as pedicel (Fig. 3E). Gena dark grayish brown (Fig. 14E). Supracervical setae 4–5 and postoculars 17–18 per side (Fig. 4E). Cibarial projections at anterolateral corners as long as width of anterior margin; medial sensilla in parallel rows as wide as sensilla campaniformia (Fig. 5E 2). Prementum and neighboring lateral membrane with 6–7 setae per side (Fig. 5E). Thorax glossy, blackish brown (Fig. 14E). Legs. Fore tarsomere I slightly shorter than II+III (Fig. 9E 1). Abdomen. Sternites dark grayish brown. Male terminalia. Cercus nearly entirely pubescent, with 32–34 setae. Female terminalia. Epigynium with approximately 7 small sensilla per side near lower posterior margin (Fig. 12D 1). Measurements (holotype /range in 5♁ and 5♀, or less if noted, paratypes, in mm): BL = 1.92/♁ 1.45–1.70, ♀ 1.65–1.88 (4♀); ThL = 0.81/♁ 0.66–0.82, ♀ 0.78–0.84; WL = 1.56/♁ 1.30–1.54, ♀ 1.46–1.61; WW = 0.74/♁ 0.60–0.72, ♀ 0.69–0.76. Indices (holotype /range in 5♁ and 5♀, or less if noted, paratypes, in ratio): FW/HW = 0.58/0.55–0.60; ch/o = 0.40/0.39–0.46; rcorb = 0.38/0.43–0.55 (4♁ and 1♀); vb = 0.26/0.32–0.43; sctl = 0.56/0.58–0.70 (3♁ and 2♀); sctlp = 1.14/1.02–1.19; C = 1.46/1.30–1.43; 4c = 1.71/1.67–2.25; 4v = 2.94/2.52–3.58; 5x = 7.14/5.07–7.56; ac = 2.98/3.12–3.63; M = 1.25/1.14–1.64. Holotype. ♁ (MZB), “ Cibodas, West Java, Indonesia, 12.xii.2004, K. T. Takano / ex inflorescence of Pinanga coronata ”. Paratypes. Indonesia: 9♁, 23♀, same data as the holotype, 14 ♁, 13♀, same data as the holotype except for 24.xii.2003 (MZB, SEHU); 10 ♁, 10♀ (#05202–21), same data as the holotype except for 24.xii.2003 (KIZ). Distribution. Indonesia (West Java). Remarks. This species resembles C. oligochaeta sp. nov., but can be distinguished from it by the diagnostic characters. Etymology. Patronym, in honor of Dr. David Grimaldi at the American Museum of Natural History, who made early contributions to the systematics of Colocasiomyia., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on page 227, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x"]}

1) Colocasiomyia pinangae Zhang & Toda, sp. nov. (Figs 2–12A, 14A) Colocasiomyia sp.1 aff. nepalensis: Sultana et al., 2006: 694; Fartyal et al., 2013: 768. Diagnosis. Surstylus with 5–7 stout, short, peg-like teeth (Fig. 11-1A 2). Pregonites apically round, with coarse serrations on ventral margin (Fig. 11-1A 4). Epigynium with 11–12 sensilla per side near lower posterior margin (Fig. 12A 1). Distal, narrow portion of hypogynial valve slightly shorter than basal, broad portion (Fig. 12A). Description (♁ and ♀). Head. Antennal pedicel blackish brown; first flagellomere approximately 1.7 times as long as pedicel; arista approximately 2.1 times as long as first flagellomere (Fig. 3A). Distance between antennal scapes narrower than scape diameter (Fig. 2A 2). Facial carina wider than 1st flagellomere (Fig. 2A 2). Supracervical setae 4–7 and postoculars 17–18 per side (Fig. 4A). Cibarial projections at anterolateral corners as long as width of anterior margin; medial sensilla 2–4 per side (Fig. 5A 2). Prementum and neighboring lateral membrane with 7–8 setae per side (Fig. 5A 2). Labellum with 10 pseudotracheae per side (Fig. 5A 1). Thorax. Two katepisternal, prominent setae longer than acrostichal setulae (Fig. 7A). Legs grayish brown, except for fore tibiae and all tarsi pale grayish yellow (Fig. 14A). Fore tarsomere I slightly longer than tarsomeres II and III combined; mid I as long as II+III+IV; hind I slightly longer than II+III+IV (Fig. 9A). Fore tarsomere II with 1 large and 1 small pegs; latter 1/3 as long as former (Fig. 9A 1). Abdomen. Tergites blackish brown (Fig. 14A). Male sternites III and IV as wide as long; V+VI distally deeply bilobed (Fig. 10A 1). Female sternites III–V as wide as long; VI wider than long; VII wider than long plate bilobed in distal half (Fig. 10A 2). Female tergite VII caudomedially deeply notched. Male terminalia. Epandrium with 2–4 long setae only on lateral to ventral portion (Fig. 11-1A 1). Cercus nearly entirely pubescent except for small, anteroventral, expanded flap, with approximately 37 setae (Fig. 11-1A 1). Phallal sheath not pubescent (Fig. 11-1A 4). Female terminalia. Perineal membrane with patch of dense, distinct warts. Measurements (holotype /range in 6♁ and 5♀ paratypes, in mm): BL (straight distance from anterior edge of pedicel to tip of abdomen) = 1.45/♁ 1.29–1.54, ♀ 1.43–1.62; ThL (distance from anterior notal margin to apex of scutellum) = 0.58/♁ 0.59–0.67, ♀ 0.60–0.67; WL (distance from humeral cross vein to wing apex) = 1.22/♁ 1.25–1.40, ♀ 1.23–1.44; WW (maximum wing width) = 0.60/♁ 0.58–0.66, ♀ 0.59–0.67. Indices (holotype /range in 6♁ and 5♀, or less if noted, paratypes, in ratio): FW/HW (frontal width / head width) = 0.55/0.53–0.60; ch/o (maximum width of gena / maximum diameter of eye) = 0.36/0.29–0.40; prorb (proclinate orbital seta length / posterior reclinate orbital seta length) = 0.85/0.84–0.97 (2♁, 1♀); rcorb (anterior reclinate orbital seta length / posterior reclinate orbital seta length) = 0.30/0.23–0.40 (3♁, 1♀); vb (subvibrissal seta length / vibrissa length) = 0.33/0.22–0.32; dcl (anterior dorsocentral seta length / posterior dorsocentral seta length) = 0.46/0.44–0.71 (4♁, 4♀); presctl (prescutellar acrostichal seta length /posterior dorsocentral seta length) = 0.42/0.25–0.40 (6♁, 4♀); sctl (basal scutellar seta length / apical scutellar seta length) = 0.54/0.54–0.63 (6♁, 4♀); sterno (anterior katepisternal seta length / posterior katepisternal seta length) = 1.00/0.94–1.26; orbito (distance between proclinate and posterior reclinate orbital setae / distance between inner vertical and posterior reclinate orbital setae) = 0.35/0.32–0.51; dcp (distance between ipsilateral dorsocentral setae / distance between anterior dorsocentral setae) = 0.50/0.57–0.65; sctlp (distance between ipsilateral scutellar setae / distance between apical scutellar setae) = 1.07/0.86–1.03; C (2nd costal section between subcostal break and R 2 +3 / 3rd costal section between R 2 +3 and R 4 +5) = 2.04/1.70–2.16; 4c (3rd costal section between R 2 +3 and R 4 +5 / M 1 between r-m and dm-m) = 1.46/1.30–1.54; 4v (M 1 between dm-m and wing margin / M 1 between r-m and dm-m) = 2.86/2.48–2.78; 5x (M 4 between dm-m and wing margin / dm-m between M 1 and M 4) = 3.18/2.50–3.35; ac (3rd costal section between R 2 +3 and R 4 +5 / distance between distal ends of R 4 +5 and M 1) = 2.63/2.24–3.04; M (M 4 between dm-m and wing margin / M 1 between r-m and dm-m) = 1.11/0.87–1.08; C3F (length of heavy setation in 3rd costal section / length of 3rd costal section) = 0.37/0.37–0.49. Holotype. ♁ (MZB), “ Cibodas, West Java, Indonesia, 12.xii.2004, K. T. Takano / ex inflorescence of Pinanga coronata ”. Paratypes. Indonesia: 24♁, 26♀, same data as the holotype (MZB, SEHU); 10 ♁, 10♀ (#05160–79), same data as the holotype except for 24.xii.2003 (KIZ). Distribution. Indonesia (West Java). Remarks. This species resembles C. nepalensis, but can be distinguished from it in having the shorter arista (in C. nepalensis: “2.5× the length of flagellomere I”; Grimaldi 1991), cibarial medial sensilla in parallel rows as wide as sensilla campaniformia (narrower than the latter; “ Fig. 7 ” in Grimaldi 1991), two equally long katepisternal setae (“Katepisternum … with 1 large seta”), and two pegs much different in size on the fore tarsomere II (“Tarsal segment on foreleg … bearing a pair of black, spine-like setae; medial one slightly longer”). Etymology. Referring to the host-plant genus Pinanga.

4) Colocasiomyia oligochaeta Zhang & Toda, sp. nov. (Figs 2–12D, 14D) Colocasiomyia sp.2 aff. nepalensis: Sultana et al., 2006: 694; Fartyal et al., 2013: 768. Diagnosis. Antennal arista approximately 1.8 times as long as first flagellomere (Fig. 3D). Distance between antennal scapes as wide as scape diameter (Fig. 2D 2). Acrostichal setulae in 6 rows (Fig. 6D). All legs entirely grayish yellow to yellow except for grayish brown tarsomere Vs (Fig. 14D). Cercus slightly projected at apicoventral corner (Fig. 11-2D). Long, tube-like, membranous aedeagus ornamented with relatively sparse and weak spinules (Fig. 11-2D). Hypogynial valves with dorsal flaps fused into small, ligulate plate at distal end of basal portion (Fig. 12D). Description (♁ and ♀). Head. Antennal, first flagellomere approximately 1.7 times as long as pedicel (Fig. 3D). Gena grayish yellow (Fig. 14D). Supracervical setae 6–9 and postoculars 13–15 per side (Fig. 4D). Cibarial projections at anterolateral corners shorter than width of anterior margin; medial sensilla in parallel rows wider than sensilla campaniformia (Fig. 5D 2). Prementum and neighboring lateral membrane with approximately 5 setae per side (Fig. 5D). Thorax. Scutum and scutellum glossy, blackish brown; pleura grayish brown (Fig. 14D). Legs. Fore tarsomere I as long as II+III (Fig. 9D 1). Abdomen. Sternites grayish brown. Male terminalia. Cercus pubescent except for anterior to ventral margin, with approximately 32 setae. Female terminalia. Epigynium with 5–6 sensilla per side near lower posterior margin (Fig. 12D 1). Measurements (holotype /range in 5♁ and 5♀, or less if noted, paratypes, in mm): BL = 1.51/♁ 1.30–1.62, ♀ 1.42–1.64 (4♀); ThL = 0.69/♁ 0.59–0.75, ♀ 0.59–0.71; WL = 1.35/♁ 1.20–1.44, ♀ 1.14–1.38; WW = 0.66/♁ 0.62–0.76, ♀ 0.54–0.72. Indices (holotype /range in 5♁ and 5♀, or less if noted, paratypes, in ratio): FW/HW = 0.56/0.56–0.62; ch/o = 0.34/0.34–0.39; rcorb = 0.44/0.47–0.60 (4♁ and 3♀); vb = 0.33/0.27–0.38; sctl = 0.62/0.61–0.74 (4♁ and 5♀); sctlp = 1.10/0.89–1.17; C = 1.44/1.28–1.47; 4c = 1.74/1.69–2.13; 4v = 2.57/2.60–3.15; 5x = 4.61/4.02–5.11; ac = 3.05/2.72–3.57; M = 1.15/1.10–1.34. Holotype. ♁ (MZB), “ Cibodas, West Java, Indonesia, 12.xii.2004, K. T. Takano / ex inflorescence of Pinanga coronata ”. Paratypes. Indonesia: 5♁, 10♀, same data as the holotype, 7 ♁, 20♀, same data as the holotype except for 24.xii.2003 (MZB, SEHU); 10 ♁, 10♀ (#05180–99), same data as the holotype except for 24.xii.2003 (KIZ). Distribution. Indonesia (West Java). Etymology. Referring to the lack of some setae from the head and thorax.

2) Colocasiomyia besaris Yafuso & Toda, sp. nov. (Figs 2–12B, 14B) Colocasiomyia sp.3 aff. nepalensis: Sultana et al., 2006: 694. Diagnosis. Surstylus with 8–10 stout, longer, peg-like teeth; dorsalmost one thickest (Fig. 11-1B 2). Pregonites apically pointed, smooth on ventral margin (Fig. 11-1B 4). Epigynium with 26–28 small sensilla per side near lower posterior margin (Fig. 12B 1). Distal, narrow portion of hypogynial valve longer than basal, broad portion (Fig. 12B). Description (♁ and ♀). Head. Antennal pedicel blackish brown; first flagellomere approximately 1.6 times as long as pedicel; arista approximately 2.1 times as long as first flagellomere (Fig. 3B). Distance between antennal scapes as wide as scape diameter (Fig. 2B 2). Facial carina wider than 1st flagellomere (Fig. 2B 2). Supracervical setae 8–10 and postoculars 18–20 per side (Fig. 4A). Cibarial projections at anterolateral corners as long as width of anterior margin; medial sensilla approximately 2–3 per side (Fig. 5B 2). Prementum and neighboring lateral membrane with 9–10 setae per side (Fig. 5B 2). Labellum with 11 pseudotracheae per side (Fig. 5B 1). Thorax. Two katepisternal, prominent setae longer than acrostichal setulae (Fig. 7B). Legs grayish brown, except for all tarsi pale grayish yellow (Fig. 14B). Fore tarsomere I slightly longer than tarsomeres II and III combined; mid I as long as II+III+IV; hind I slightly longer than II+III+IV (Fig. 9B). Fore tarsomere II with 1 large and 1 small pegs; latter 1/3 as long as former (Fig. 9B 1). Abdomen. Tergites blackish brown (Fig. 14B). Male sternites III and IV as wide as long; V+VI distally deeply bilobed (Fig. 10B 1). Female sternites III–V as wide as long; VI wider than long; VII wider than long plate bilobed in distal half (Fig. 10B 2). Female tergite VII mid-dorsally separated into 2 lateral plates. Male terminalia. Epandrium with 2–4 long setae only on lateral to ventral portion (Fig. 11-1B 1). Cercus nearly entirely pubescent except for roundish, ventral flap, with approximately 40 setae (Fig. 11-1B 1). Phallal sheath partly pubescent (Fig. 11-1B 4). Female terminalia. Perineal membrane with patch of dense, distinct warts. Measurements (holotype /range in 2♁ and 1♀ paratypes, in mm): BL = 1.51/♁ 1.35–1.55, ♀ 1.70; ThL = 0.67/♁ 0.72–0.73, ♀ 0.72; WL = 1.39/♁ 1.46–1.55, ♀ 1.47; WW = 0.66/♁ 0.72, ♀ 0.72. Indices (holotype /range in 2♁ and 1♀, or less if noted, paratypes, in ratio): FW/HW = 0.54/0.53–0.56; ch/o = 0.38/0.32–0.40; prorb = 0.83/0.82–0.97; rcorb = 0.30/0.25–0.32; vb = 0.21/0.24–0.29; dcl = 0.51/0.52–0.54 (1♁ and 1♀); presctl = 0.30/0.35–0.41; sctl = 0.75/0.71–0.73 (1♁ and 1♀); sterno = 1.07/1.28–1.42; orbito = 0.35/0.38–0.45; dcp = 0.58/0.68–0.77; sctlp = 0.81/0.91–1.07; C = 2.10/1.82–2.18; 4c = 1.32/1.22–1.39; 4v = 2.54/2.33–2.61; 5x = 2.78/1.82–3.23; ac = 2.50/2.36–2.87; M = 0.99/0.91–1.02; C3F = 0.41/0.46–0.54. Holotype. ♁ (MZB), “ Cibodas, West Java, Indonesia, 12.xii.2004, K. T. Takano / ex inflorescence of Pinanga coronata ”. Paratypes. Indonesia: 12♁, 4♀, same data as the holotype (MZB, SEHU); 2♁ (#05200–01), 1♀ (#05226), same data as the holotype except for 24.xii.2003 (KIZ). Distribution. Indonesia (West Java). Remarks. This species resembles C. pinangae sp. nov., but can be distinguished from it by the diagnostic characters. Etymology. The specific name derives from “besar” (meaning “large”) of Indonesian, referring to its larger body size among the members of the zeylanica group., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on pages 223-224, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x"]}

3) Colocasiomyia luciphila Zhang & Toda, sp. nov. (Figs 2–12C, 14C) Colocasiomyia sp.5 aff. nepalensis: Fartyal et al., 2013: 769. Diagnosis. Antennal, first flagellomere approximately 2.4 times as long as pedicel; arista approximately 1.6 times as long as first flagellomere (Fig. 3C). Fore tarsomere II with 2 pegs; smaller peg slightly longer than 1/2 of larger one (Fig. 9C 1). Male sternites III and IV longer than wide; V+VI concave on distal margin (Fig. 10C 1). Female sternites III–VI longer than wide; VII much wider than long, slightly convex on distal margin (Fig. 10C 2). Surstylus with 6–7 stout, longer, peg-like teeth nearly uniform in thickness (Fig. 11-1C 1,2). Pregonites apically roundish, smooth on ventral margin (Fig. 11-1C 4a). Epigynium with 2–6 small sensilla per side near lower posterior margin (Fig. 12C 1). Distal, narrow portion of hypogynial valve half as long as basal, broad portion (Fig. 12C). Description (♁ and ♀). Head. Antennal pedicel grayish brown. Distance between antennal scapes as wide as scape diameter (Fig. 2C 2). Facial carina narrower than 1st flagellomere (Fig. 2C 2). Supracervical setae 6–8 and postoculars 13–18 per side (Fig. 4C). Cibarial projections at anterolateral corners slightly longer than width of anterior margin; medial sensilla approximately 3–4 per side (Fig. 5C 2). Prementum and neighboring lateral membrane with 7–8 setae per side (Fig. 5C 2). Labellum with 11–13 pseudotracheae per side (Fig. 5C 1). Thorax. Prescutellar, acrostichal setae only slightly longer than acrostichal setulae (Fig. 6C). Only one prominent katepisternal seta longer than acrostichal setulae (Fig. 7C). Legs grayish brown, except for all tarsi pale grayish yellow (Fig. 14C). Fore, mid and hind tarsomere Is as long as II+III, II+III+IV and II+III+IV+V, respectively (Fig. 9C). Abdomen. Tergites dark grayish brown (Fig. 14C). Female tergite VII mid-dorsally separated into 2 lateral plates. Male terminalia. Epandrium with approximately 4 long setae only on lateral to ventral portion (Fig. 11-1C 1). Cercus entirely pubescent, round on ventral margin, with 28–29 setae (Fig. 11-1C 1). Phallal sheath partly pubescent (Fig. 11-1C 4,b). Female terminalia. Perineal membrane without patch of dense, distinct warts. Measurements (holotype /range in 1♁ and 3♀ paratypes, in mm): BL = 1.51/♁ 1.59, ♀ 1.40–1.92; ThL = 0.60/♁ 0.60, ♀ 0.48–0.75; WL = 1.22/♁ 1.23, ♀ 1.06–1.55; WW = 0.57/♁ 0.61, ♀ 0.50–0.73. Indices (holotype /range in 1♁ and 3♀, or less if noted, paratypes, in ratio): FW/HW = 0.48/0.49–0.55; ch/o = 0.29/0.33–0.38; prorb = 0.86/0.82–1.08; rcorb = 0.32/0.29–0.40; vb = 0.17/0.18–0.25 (3♀); dcl = 0.39/0.35– 0.40 (1♁ and 2♀); presctl = 0.20/0.21–0.25 (1♁ and 2♀); sctl = 0.56/0.52–0.60; orbito = 0.45/0.33–0.47; dcp = 0.54/0.49–0.59; sctlp = 0.93/0.88–1.03; C = 1.57/1.49–1.64; 4c = 1.61/1.39–1.64; 4v = 2.39/2.20–2.63; 5x = 2.61/2.62–2.75; ac = 3.27/2.49–3.09; M = 0.90/0.78–0.92; C3F = 0.60/0.49–0.72. Holotype. ♁ (RDID), “ Lambir Hills National Park, Sarawak, Malaysia, 25–26.i.2012, ex light trap (LT-35), M.J. Toda ”. Paratypes. Malaysia: 5♀, same data as the holotype (SEHU); 1♀, same except for 26–27.i.2012, 1♀, same except for 23.i.2012, ex florescent light, 1♀, same except for 25.i.2012, ex florescent light (RDID); 1♁ (#05225), same data as the holotype, 1♀ (#05223), same except for 23–24.i.2012 (Lot #229), 1♀ (#05224), same except for 24–25.i.2012 (Lot #230) (KIZ). Distribution. Malaysia (Sarawak). Remarks. This species resembles C. nepalensis in having only one prominent, katepisternal seta and two pegs less different in length on the fore tarsomere II, but can be distinguished from it by the arista 1.6 times as long as first flagellomere (in C. nepalensis: “2.5× the length of flagellomere I”; Grimaldi 1991), cibarial medial sensilla in parallel rows as wide as sensilla campaniformia (narrower than the latter; “ Fig. 7 ” in Grimaldi 1991), the facial carina narrower than 1st flagellomere (wider than the latter; “ Fig. 6 ” in Grimaldi 1991), and the absence of membranous aedeagus (“Distiphallus fusiform, apically with fine spicules, covered with membrane”). On the other hand, the following characters suggest some relatedness to the oligochaeta species subgroup described below: only one prominent, katepisternal seta; prescutellar, acrostichal setae less differentiated; and male sternites III and IV and female III–VI longer than wide. Etymology. Referring to its nature of being attracted to light., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on pages 224-225, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["Fartyal, R. S., Gao, J. J., Toda, M. J., Hu, Y. G., Takano, K. T., Suwito, A., Katoh, T., Takigahira, T. & Yin, J. T. (2013) Colocasiomyia (Diptera: Drosophilidae) revised phylogenetically, with a new species group having peculiar lifecycles on monsteroid (Araceae) host plants. Systematic Entomology, 38, 763 - 782. https: // doi. org / 10.1111 / syen. 12027","Grimaldi, D. A. (1991) Systematics of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae): cladistics, a new generic synonym, new record, and a new species from Nepal. Entomologica Scandinavica, 22, 417 - 426. https: // doi. org / 10.1163 / 187631291 X 00219"]}

Colocasiomyia zeylanica species group Colocasiomyia zeylanica species group: Sultana et al., 2006: 694; Fartyal et al., 2013: 769. Diagnosis (modified from Fartyal et al. 2013). Pedicel with uniformly short, stout setae only, lacking prominently long one(s) (ch.2-1; Fig. 3, “ Fig. 6 ” in Grimaldi 1991). Two peg-like setae present only on fore tarsomere II (ch.36- 0, ch.38-0; Fig. 9A–E 1, “ Fig. 1C ” in Okada 1986, “ Fig. 9 ” in Grimaldi 1991). Hypandrium forming like tube surrounding phallus (ch.66-2; Fig. 11). Gonocoxites absent (ch.69-4; Fig. 11). Shared characters. Distance between antennal scapes wider than half of scape diameter (ch.1-1; Fig. 2A–E 2, “ Fig. 1A ” in Okada 1986, “ Fig. 6 ” in Grimaldi 1991). Antennal, first flagellomere grayish yellow, with small invaginated organ but no hollow organ (ch.3-0; Fig. 3, “ Fig. 6 ” in Grimaldi 1991); arista entirely densely pubescent, without branches (ch.5-1; Fig. 3, “ Fig. 1A, B ” in Okada 1986, “ Fig. 6 ” in Grimaldi 1991). Eye dark red (Fig. 14). Frons blackish brown; frontal vittae mat. Supracervical setae distally more or less curved, longer than inner, occipital setae (ch.11-0, ch.12-0; Fig. 4). Palpus pale grayish yellow, flat or convex in ventrodistal portion (ch.16-0; Fig. 5, “ Fig. 1B ” in Okada 1986, “ Fig. 7 ” in Grimaldi 1991). All cibarial sensilla very small or minute; anterior sensilla irregularly arranged in square or triangle (ch.18-1, ch.19-1; Fig. 5A–E 2, “ Fig. 7 ” in Grimaldi 1991). Proboscis without a pair of lateral, spherical bumps covered with numerous, short, stout setae (ch.22-0; Fig. 5, “ Fig. 7 ” in Grimaldi 1991). Additional, presutural, dorsocentral setae absent (Fig. 6, “ Fig. 1A ” in Okada 1986). Basal and apical, scutellar setae nearly parallel (Fig. 6). Halter grayish brown. Wing hyaline; veins yellow (Fig. 8). Fore tarsomere II slightly elongated below (ch.34-1; Fig. 9A–E 1, “ Fig. 1C ” in Okada 1986, “ Fig. 9 ” in Grimaldi 1991). Preapical setae absent on all tibiae (ch.41-1; Fig. 9, “ Fig. 1A ” in Okada 1986). Dorsomedian, terminal, down-curved seta on tarsomere Vs not longer than claw (ch.42-0; Fig. 9). Mid tibia apicoventrally with 2 (1 long and 1 small) stout setae (ch.39-0; Fig. 9A–E 2). Spiracles simple, small (ch.44-0). Male sternite VI pubescent entirely, without process (ch.48-0, ch.50-0; Fig. 10A–E 1). Epandrium anteroventrally much elongated, not fused to cercus, without peg-like, stout seta at posteroventral apex; apodeme narrow lobe on anteromedial to -dorsal margin or absent (ch.55- 1, ch.57-0, ch.58-0, ch.62-0; Figs 11-1A–C 1,2, 11-2D, E). Hypandrial setae absent (ch.67-1; Figs 11-1A–C 3,4, 11-2D, E). Postgonites fused to aedeagal sheath, forming together phallal-sheath tube distally more or less separated into a pair of lateral lobes (ch.71-1, ch.72-1, ch.73-0; Figs 11-1A–C 3,4, 11-2D, E). Phallapodeme mostly rod-like (ch.77-0; Figs 11-1A–C 3,4, 11-2D, E; “ Fig.10 ” in Grimaldi 1991). Phallal guide absent (ch.78-0; Figs 11-1A–C 4, 11-2D, E). Epiproct nearly entirely pubescent (ch.79-0; Fig. 12A–E 1). Epigynium pubescent only on dorsal portion (ch.82-0; Fig. 12A–E 1). Hypogynial, anteroventral bridge absent (ch.85-1; Fig. 12A–E 2); valves separated from each other, distally differentiated into more or less narrow process from basal, broad portion, with only trichoid sensilla (ch.86- 0, ch.87-1, ch.88-1; Fig. 12). Included taxa. Colocasiomyia zeylanica species subgroup; and C. oligochaeta subgroup. Key. The digital, multiple-entry key “ Colocasiomyia zeylanica species group” to all species of this group has been constructed partly based on the character matrix of Appendix 2, and is available from the following website: https://evolgen.biol.se.tmu.ac.jp/Classification/index.jsp (Biological Classification and Identification System)., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on page 221, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x","Fartyal, R. S., Gao, J. J., Toda, M. J., Hu, Y. G., Takano, K. T., Suwito, A., Katoh, T., Takigahira, T. & Yin, J. T. (2013) Colocasiomyia (Diptera: Drosophilidae) revised phylogenetically, with a new species group having peculiar lifecycles on monsteroid (Araceae) host plants. Systematic Entomology, 38, 763 - 782. https: // doi. org / 10.1111 / syen. 12027","Grimaldi, D. A. (1991) Systematics of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae): cladistics, a new generic synonym, new record, and a new species from Nepal. Entomologica Scandinavica, 22, 417 - 426. https: // doi. org / 10.1163 / 187631291 X 00219","Okada, T. (1986) Taximetrical analyses of costal chaetotaxy of the genus Drosophilella Duda, with description of a new species from Sri Lanka (Diptera, Drosophilidae). Proceedings of the Japanese Society of Systematic Zoology, 34, 53 - 59."]}

Genus Colocasiomyia de Meijere, 1914 Colocasiomyia de Meijere, 1914: 272 (in Borboridae). Type species: Colocasiomyia cristata de Meijere, 1914. Colocasiomyia: Duda, 1924: 177; Okada, 1988: 34; Grimaldi, 1991: 417; Sultana et al., 2006: 693; Takano et al., 2021: 19. Platyborborus de Meijere, 1914: 273 (in Sphaeroceridae). Type species: Platyborborus crassipes de Meijere, 1914. Syn. Grimaldi, 1991: 419. Drosophilella Duda, 1923: 25. Type species: Drosophilella seminigra Duda, 1923. Syn. Okada, 1988: 34. Diagnosis (modified from Takano et al. 2021). Fore tarsus with heavily sclerotized peg-like setae in both sexes (ch.35-1). Sclerotized spermathecal capsule absent (ch.89-1)., Published as part of Zhang, Guang, Gao, Jian-Jun, Takano, Kohei Takenaka, Yafuso, Masako, Suwito, Awit, Meleng, Paulus Ak & Toda, Masanori J., 2023, Phylogenetic classification and palm-inflorescence anthophily of the Colocasiomyia zeylanica species group (Diptera: Drosophilidae), with descriptions of five new species, pp. 201-238 in Zootaxa 5278 (2) on page 221, DOI: 10.11646/zootaxa.5278.2.1, http://zenodo.org/record/7905995, {"references":["de Meijere, J. C. H. (1914) Studien ¸ ber s ¸ dostasiatische Dipteren IX. Tijdschrift voor Entomologie, 57, 137 - 275.","Duda, O. (1924) Beitrag zur Systematik der Drosophiliden unter besonderer Ber ¸ cksichtigung der palaarktischen u. orientalischen Arten (Dipteren). Wiegmann's Archiv fur Naturgeschichte, 90 (A) 3, 172 - 234, 7 pls.","Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34 - 39.","Grimaldi, D. A. (1991) Systematics of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae): cladistics, a new generic synonym, new record, and a new species from Nepal. Entomologica Scandinavica, 22, 417 - 426. https: // doi. org / 10.1163 / 187631291 X 00219","Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x","Takano, K. T., Gao, J. - J., Hu, Y. - G., Li, N. - N., Yafuso, M., Suwito, A., Repin, R., Pungga, R. A. S., Meleng, P. A., Kaliang, C. H., Chong, L. & Toda, M. J. (2021) Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species. Zootaxa, 5079 (1), 1 - 70. https: // doi. org / 10.11646 / zootaxa. 5079.1.1","Duda, O. (1923) Die orientalischen und australischen Drosophiliden-Arten (Dipteren) des Ungarischen National-Museums zu Budapest. Annales Historico-Naturales Musei Nationalis Hungarici, 20, 24 - 59."]}

A list of ants of the subfamilies Leptanillinae, Proceratiinae, Amblyoponinae, Ponerinae, Dorylinae, Dolichoderinae, Ectatomminae and Formicinae from Lambir Hills National Park, Sarawak, Borneo is presented with biological information. In this report 347 species in 61 genera are recorded, with 10 species newly recorded from Borneo. Literature treating species of the abovementioned subfamilies from Lambir Hills National Park is listed. In total (Part I & II), 579 species in 96 genera are recorded from this national park, representing the largest species number so far known in one place in Asia. Among the 579 species only 53% (307) are identified to species, the others no doubt including many species new to Borneo or even to science.

To assess the diversity of vascular epiphytes in primary lowland mixed dipterocarp forest in Borneo, field surveys were conducted in Lambir Hills National Park, Sarawak, Malaysia in 2014–2019. A total of 183 species including 72 species of eudicots (2 species were unknown at family level) (39 %), 71 monocots (39 %), 37 pteridophytes (20 %) and 3 lycophytes (2 %), are recorded. A checklist, with voucher specimens and notes on their identification and ecological information for each species, are provided. Dapania racemosa Korth. (Oxalidaceae) formerly recognized as a liana, was observed as an epiphyte for the first time.

6) Colocasiomyia kinabaluana Toda & Takano, sp. nov. (Figs 9J, 15F, 21) Colocasiomyia sp.2 aff. sulawesiana: Sultana et al., 2006: 694; Toda & Lakim, 2011: 264; Takano et al., 2011: 22; Takano et al., 2012: 559, Supplemental File 1; Fartyal et al., 2013, Fig. 6. Diagnosis. A pair of processes on male abdominal sternite VI short ( Description (♂ and ♀). Head. Supracervical setae 6–9 per side, distally more or less curved, longer than inner occipital setae. Cibarial, medial sensilla approximately 3 per side; posterior sensillum 1 per side. Supralateral setae outside prementum 3–4 per side. Thorax. Anterior dorsocentral setae just beside transverse suture. Abdomen. Male sternite III and IV longer than wide (Fig. 21A). Female sternites V and VI longer than wide (Fig. 21B). Male terminalia. Epandrium pubescent except for anterior margin, anteroventral elongation and ventral portion, with 1–2 setae on dorsal portion and 21–23 setae thicker than cercal setae on ventral lobe of each side (Fig. 21C). Cercus pubescent on dorsal 2/3 but not on anterior margin and ventral portion, with 38–39 setae, extended below, apically forming small lobe marginally fringed with small setae (Fig. 21C). Phallapodeme less sclerotized thin plate, nearly perpendicular to phallal axis (Fig. 21D). Female terminalia. Hypoproct not pubescent. Oviscapt longer than phallus (apodeme + sheath), with 21–22 ovisensilla only on distal portion occupying less than 1/2 length of oviscapt (Fig. 21G). Indices (range of 10♂ and 10♀): FW/HW = 0.57–0.67, ch/o = 0.37–0.53, prorb = 0.94–1.15, rcorb = 0.25–0.54, vb = 0.23–0.38, dcl = 0.58–0.86, sctl = 0.68–0.86, sterno = 0.83–1.02, orbito = 0.44–0.84, dcp = 1.02–1.33, sctlp = 1.07–1.36, C = 1.68–2.38, 4c = 1.02–1.37, 4v = 1.45–1.86, 5x = 0.72–1.16, ac = 2.18–3.01, M = 0.35–0.48. Puparium (3rd instar larva). Segments with stout spicules on ventral surface; anterior spiracle sessile, with a bundle of approximately 4 short branches; caudal segments elongate, with many small spicules, ending in a Vshaped pair of posterior spiracles (Fig. 21H,I). Mouth hook less expanded medioventrally in lateral view; distal blade as long as basal portion, apically pointed, weakly curved downward, with two rows of small, acute teeth on submedial to subapical portion of ventral margin (Fig. 21J). Holotype. ♂ (BORN), “ Kota Kinabalu, Sabah, Malaysia, 3.i.1999, ex Alocasia macrorrhizos, M.J. Toda ”. Paratypes. Malaysia: 23♂, 10♀, same data as the holotype; 6♂, ditto except 12.viii.2004, K. T. Takano leg. (BORN, SEHU). Philippines: 7♂, 6♀, Legaspi, S. Luzon, 13°08’52”N 123°44’04”E, 30.xi.2012, ex A. macrorrhizos, P.J. Matthews, M. Medecilo & J.R. Castillo leg.; 10♂, 10♀, Gandara, Samar, 12°00'56"N 124°47'43"E, 2.xii.2012, ex A. macrorrhizos, P.J. Matthews, M. Medecilo & J.R. Castillo leg.; 2♂, Tanuan, N. Leyte, 11°07'03"N 125°00'55"E, 3.xii.2012, ex A. macrorrhizos, P.J. Matthews, M. Medecilo & J.R. Castillo leg.; 10♂, 10♀, Sabang, Palawan, 10°11'24"N 118°54'00"E, 5.ii.2013, ex A. macrorrhizos, P.J. Matthews, E.M. Agoo & D.A. Madulid leg. (MPMP, SEHU). Distribution. Borneo (Sabah), Philippines (Luzon, Samar, Leyte, Palawan). Remarks. This species is coupled with C. sulawesiana in the molecular phylogenetic tree (BP = 87; Fig. 2) and the cladogram (BP = 100; Fig. 13), and shares the oviscapt longer than the phallus (apodeme + sheath) as a synapomorphy (ch.49-1) with it. However, they can be distinguished from each other by their diagnostic characters. Etymology. Referring to the type locality.

9) Colocasiomyia sabahana Toda, sp. nov. (Figs 9L, 11F,G, 15I, 24C,E,G,K,M,O,Q) Colocasiomyia sp.4 aff. diconica: Sultana et al., 2006: 694, Toda & Lakim, 2011: 264; Takano et al., 2011: 26. Diagnosis. Additional dorsocentral setae on presutural area absent.A pair of processes on male abdominal sternite VI slightly curved inward (Fig. 9L). Epandrium triangularly angled at posteroventral corner; anteroventral elongation as broad as cercus in lateral view (Fig. 24G). Female abdominal sternite VI wider than long (Fig. 24E). Oviscapt broader than hypoproct, apically roundish in lateral view (Fig. 24K). Description (♂ and ♀). Head. Supralateral setae outside prementum approximately 2 per side. Thorax. Basal scutellar setae short, not reaching to half of apical scutellar setae. Legs. Mid tibia with 2 apical, stout setae. Abdomen. Female sternite IV as long as wide (Fig. 24E). Male terminalia. Epandrium with approximately 1 seta on lateral portion and 14–16 setae on posteroventral portion (Fig. 24G). Median piece of subepandrial sclerite medially and anteriorly bilobed into long lobes; lateral pieces long, narrow. Cercus with 45–48 setae, extended below, apically forming small lobe with many small setae (Fig. 24G). Female terminalia. Oviscapt with 14–15 ovisensilla (Fig. 24K). Indices (range of 10♂ and 10♀): FW/HW = 0.55–0.65, ch/o = 0.34–0.45, prorb = 0.83–1.04, rcorb = 0.22–0.47, vb = 0.25–0.39, dcl = (n/a), sctl = 0.51–0.77, sterno = 0.56–0.83, orbito = 0.48–0.77, dcp = 0.82–1.21, sctlp = 1.08–1.64, C = 1.62–2.56, 4c = 0.83–1.38, 4v = 1.31–1.94, 5x = 0.74–1.25, ac = 1.98–3.74, M = 0.25–0.56. Holotype. ♂ (KPSP), “ Park Headquarters, Mt. Kinabalu, Sabah, Malaysia, 10.viii.2003, ex Alocasia scabriuscula, M.J. Toda ”. Paratypes. Malaysia: 11♂, 11♀, same data as the holotype (KPSP, SEHU). Distribution. Borneo (Sabah). Etymology. Referring to the type locality., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 38-39, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x","Toda, M. J. & Lakim, M. B. (2011) Genus Colocasiomyia (Drosophilidae: Diptera) in Sabah, Bornean Malaysia: high species diversity and use of host aroid inflorescences. Entomological Science, 14, 262 - 270. https: // doi. org / 10.1111 / j. 1479 - 8298.2011.00452. x","Takano, T. K., Suwito, A., Gao, J. J. & Yin, J. T. (2011) Molecular phylogeny of the cristata species group of the genus Colocasiomyia (Diptera: Drosophilidae). Low Temperature Science, 69, 19 - 28."]}

2) Colocasiomyia grandis Toda, sp. nov. (Figs 5B, 9E, 15B, 17) Colocasiomyia sp. aff. stamenicola: Sultana et al., 2006: 694; Takano et al., 2011: 22; Fartyal et al., 2013: 768. Diagnosis. Patch covered with only minute pubescence present on anterodorsal portion of fore tarsomere I (ch.13-0; Fig. 17A). Male abdominal sternite VI with single, large process bearing neither prominent seta nor sclerotized claw on apex but small setae on submedial to distal portion (Figs 9E, 17B). Description (♂ and ♀; not repeating characters common to C. ecornuta sp. nov.). Head. Supracervical setae 5–6 per side, distally more or less curved, longer than inner occipital setae. Anterior reclinate orbital seta small, situated slightly behind proclinate orbital seta. Eye with stout, dense interfacetal setulae. Distance between antennal sockets wider than half of socket width. Cibarial, medial sensilla (3–4 per side) in parallel rows as wide as sensilla campaniformia; posterior sensillum 1 per side. Supralateral seta outside prementum 1 per side. Thorax. Anterior dorsocentral setae far behind transverse suture (Fig. 5B). Prescutellar acrostichal setae present (Fig. 5B). Katepisternal setae large; posterior one longer than longest, postpronotal seta. Apical scutellar setae nearer to each other than to basal scutellar seta (Fig. 5B). Wing. Costal setae in middle row all apically blunt, heavy, peg-like. Abdomen. Male: sternites III and IV wider than long, medially notched on posterior margin; V medially concaved on posterior margin (Fig. 17B). Female: sternites IV and V medially concaved on posterior margin; VI longer than wide, deeply bilobed into lateral, narrow plates (Fig. 17C). Male terminalia. Epandrium pubescent except for ventral portion, posteroventrally extended as ventral lobe pointed apically, with 16–19 setae as thick as cercal setae on ventral portion of each side but none on lateral to dorsal portion (Fig. 17D). Surstylus long, narrow plate with 4 somewhat trichoid, recurved setae apically, 1 nearly straight seta apicoventrally and 1 minute seta subapically (Fig. 17D). Median piece of subepandrial sclerite broad, somewhat quadrate plate; lateral pieces long, narrow. Cercal dorsal half pubescent and densely setigerous; ventral portion not pubescent, extended below, with small setae sparsely on surface and tiny setae densely on apical margin (Fig. 17D). Phallal sheath not pubescent, as thick as wide, apically somewhat hooked in lateral view, not projected at dorsobasal corner in lateral view; phallapodeme sclerotized plate directed nearly along phallal axis (Fig. 17E). Gonocoxites long, narrow slips (Fig. 17E). Female terminalia. Epiproct and hypoproct pubescent (Fig. 17F). Oviscapt subapically widest, apically round, with 14–16 ovisensilla only on distal portion occupying less than 1/2 length of oviscapt (Fig. 17F). Indices (range of 10♂ and 10♀): FW/HW = 0.50–0.72, ch/o = 0.41–0.56, prorb = 0.77–1.09, rcorb = 0.24–0.50, vb = 0.33–0.56, dcl = 0.50–0.67, sctl = 0.46–0.88, sterno = 0.42–0.78, orbito = 0.43–0.75, dcp = 0.64–0.87, sctlp = 0.98–1.26, C = 1.81–2.52, 4c = 0.85–1.20, 4v = 1.44–1.83, 5x = 0.77–1.36, ac = 2.43–2.96, M = 0.32–0.51, C3F [length of heavy setation in 3rd costal section / (length of heavy setation in 3rd costal section + length of light setation in 3rd costal section)] = 0.30–0.43. Puparium (3rd instar larva). Segments with stout spicules on ventral surface; anterior spiracle with stalk ending in a whorl of 11–12 branches; stalk as long as longest branch; caudal segments not elongate, with many small spicules, ending in a pair of short posterior spiracles (Fig. 17G–I). Mouth hook strongly, triangularly expanded medioventrally in lateral view; distal blade strongly curved downward (Fig. 17J). Holotype. ♂ (KIZ), “ Menglun, Xishuangbanna, Yunnan, China, 940 m a.s.l., 10.iii.2003, ex Alocasia odora (Roxburgh) K. Koch, M.J. Toda ”. Paratypes. China: 10♂, 10♀, same data as the holotype; 5♂, 5♀, ditto except 22.v.2019, S. Ling leg.; 5♂, 5♀, ditto except 17.iv.2017, ex Colocasia esculenta (L.) Schott; 2♂, 4♀, ditto except 7.v.2018, ex Leucocasia gigantea (Blume) Schott; 1♂ (#10149), 2♀ (#10495, #10496), Tongbiguan, Yingjiang, Yunnan, 1300 m a.s.l., 5.vi.2019, ex Alocasia yunqianum Ma, Li & Yin, Z. Ma leg. (KIZ, SEHU). Other specimens examined. Vietnam: 2♀, Pu Mat, Con Cuông District, Nghe An, 30.ix.2007, ex Alocasia aff. odora, M. Yafuso leg. (SEHU). Distribution. China (Yunnan), Vietnam. Remarks. This species is unique, having the single process on the male abdominal sternite VI, among species of the cristata subgroup; all the species other than C. ecornuta sp. nov. have the paired processes (ch.24-1; Fig. 9F–R). On the other hand, the morphological similarity in the process on the male sternite VI (a trace of fusion or separation of the process, and the lack of apical claw; Fig. 9D,E) and the fore tarsus (the patch covered with only minute pubescence on tarsomere I, and the 4 pegs on II; Fig. 17A) suggests the relationship between C. grandis sp. nov. and the most basal species, C. stamenicola, of the C. colocasiae subgroup. Etymology. Referring to the relatively large body size.

11) Colocasiomyia pistilicola (Carson & Okada, 1980) (Figs 9N, 25) Drosophilella pistilicola Carson & Okada, 1980: 18. Colocasiomyia pistilicola: Okada, 1988: 36. Diagnosis. A pair of processes on male abdominal sternite VI basally fused to each other, forming somewhat Vshaped plate (Fig. 9N). Supplementary description. Supracervical setae 4–5 per side. Eye with stout, dense interfacetal setulae. Cibarial, medial sensilla approximately 2 per side; posterior sensillum 1 per side. Supralateral setae outside prementum approximately 3 per side. Prescutellar acrostichal setae absent. Apical scutellar setae nearer to each other than to basal scutellar seta. Costal setae in middle row all weak, trichoid (Fig. 25A). Mid tibia with approximately 3 apical, stout setae (Fig. 25C). Epandrium pubescent except for anterior margin and anteroventral elongation, posteroventrally extended, with 0–2 setae on lateral to dorsal portion and 17–21 setae on ventral portion of each side (Fig. 25E,F). Cercus pubescent except for anterior margin and ventro-apical small lobe, with 35–39 setae (Fig. 25E,F). Oviscapt about 9 times as long as wide, with 15–16 ovisensilla (Fig. 25I). Specimens examined. Papua New Guinea: 2♂ 2♀ paratypes, Lae, 30.viii.1977, ex Colocasia esculenta, H.L. Carson and T. Okada leg. (NSMT). Distribution. Papua New Guinea., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on page 40, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Carson, H. L. & Okada, T. (1980) Drosophilidae associated with flowers in Papua New Guinea. I. Colocasia esculenta. Kontyu, 48, 15 - 29.","Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34 - 39."]}

1) Colocasiomyia ecornuta Toda & Takano, sp. nov. (Figs 5G, 9A, 15A, 16) Colocasiomyia sp.3 aff. sulawesiana: Sultana et al., 2006: 694. Diagnosis. Outer vertical seta absent (Fig. 16A). Katepisternal setae minute (Fig. 5G). Scutellum with several setulae scattered on dorsal surface (Fig. 16B). Male abdominal sternite VI without any process (Figs 9A, 16C). Cercus ventrally narrowed, strongly curved posteriad, apically forming somewhat triangular lobe bearing 1 long and many short setae (Fig. 16E). Phallal sheath pubescent on dorsal surface, gently curved ventrad, wider than thick, distally narrowing, basally roundly dilated, apically knobbed, subapically with small, acute projection on right margin (Fig. 16F–H). Gonocoxites broad, somewhat triangular plates (Fig. 16G). Neither epiproct nor hypoproct pubescent (Fig. 16I). Oviscapt distally narrowing, apically obliquely truncated and with 6–7 trichoid ovisensilla (Fig. 16J). Description (♂ and ♀). Head. Supracervical setae 7–11 per side, nearly straight, as long as inner occipital setae. Anterior reclinate orbital seta long, situated slightly anteriorly to proclinate orbital seta; additional, interfrontal setulae on fronto-orbital plate present (Fig. 16A). Eye with fine, sparse interfacetal setulae (Fig. 16A). Distance between antennal sockets narrower than half of socket width (Fig. 16A). Cibarial, medial sensilla (1–3 per side) in parallel rows narrower than sensilla campaniformia; posterior sensillum 1 per side. Supralateral setae outside prementum 3–4 per side. Thorax.Anterior dorsocentral setae just beside or slightly behind transverse suture (Fig. 16B). Neither additional dorsocentral setae on presutural area nor prescutellar acrostichal setae present; acrostichal setulae in 4 rows (Fig. 16B). Basal scutellar setae short, not reaching to half of apical scutellar setae; apical scutellar setae much nearer to each other than to basal scutellar seta (Fig. 16B). Wing. Costal setae in middle row all weak, trichoid. Legs. Patch covered with only minute pubescence absent on anterodorsal portion of fore tarsomere I. Mid tibia with 2 apical, stout setae. Abdomen. Male: sternite III longer than wide; IV as wide as long (Fig. 16C). Female: sternites III–V longer than wide; VI wider than long (Fig. 16D). Male sternite VI not pubescent at least partly (Fig. 9A). Male terminalia. Epandrium pubescent on dorsal and posterolateral portions, roundish on posteroventral margin, with 2–3 setae near posterodorsal margin and 11–12 setae thicker than cercal setae on posteroventral portion of each side, but lacking phragma (Fig. 16E). Surstylus long, somewhat triangular plate with 2 stout teeth dorsally and 3 long, recurved, trichoid setae ventrally on distal margin (Fig. 16E). Median piece of subepandrial sclerite mostly bilobed into long, highly wrinkled lobes connected only at posterior end; lateral pieces less sclerotized broad plates. Cercus nearly entirely pubescent, with approximately 22 setae, including especially long, most ventral one, on dorsal, oval portion (Fig. 16E). Phallapodeme less sclerotized thin plate, nearly perpendicular to phallal axis (Fig. 16G). Female terminalia. Oviscapt shorter than phallus (apodeme + sheath) (Fig. 16G,H,J). Indices (range of 4♂ and 4♀): FW/HW (frontal width / head width) = 0.59–0.66, ch/o (maximum width of gena / maximum diameter of eye) = 0.49–0.57, prorb (proclinate orbital seta length / posterior reclinate orbital seta length) = 0.86–0.95, rcorb (anterior reclinate orbital seta length / posterior reclinate orbital seta length) = 0.54–0.70, vb (subvibrissal seta length / vibrissa length) = 0.30–0.51, dcl (anterior dorsocentral seta length / posterior dorsocentral seta length) = 0.69–1.00, sctl (basal scutellar seta length / apical scutellar seta length) = 0.69–0.78, sterno (anterior katepisternal seta length / posterior katepisternal seta length) = 0.60–0.79, orbito (distance between proclinate and posterior reclinate orbital setae / distance between inner vertical and posterior reclinate orbital setae) = 0.42–0.58, dcp (distance between ipsilateral dorsocentral setae / distance between anterior dorsocentral setae) = 0.98–1.19, sctlp (distance between ipsilateral scutellar setae / distance between apical scutellar setae) = 2.00, C (2nd costal section between subcostal break and R 2+3 / 3rd costal section between R 2+3 and R 4+5) = 1.08–1.56, 4c (3rd costal section between R 2+3 and R 4+5 / M 1 between r-m and dm-m) = 1.16–1.51, 4v (M 1 between dm-m and wing margin / M 1 between r-m and dm-m) = 1.70–1.97, 5x (M 4 between dm-m and wing margin / dm-m between M 1 and M 4) = 1.76–2.00, ac (3rd costal section between R 2+3 and R 4+5 / distance between distal ends of R 4+5 and M 1) = 2.67–3.38, M (M 4 between dm-m and wing margin / M 1 between r-m and dm-m) = 0.49–0.59. Puparium (3rd instar larva). Segments with stout spicules on ventral surface; some lateral ones large, clawlike; anterior spiracle absent; caudal segments elongate, very narrow, without spicules, ending in a pair of minute posterior spiracles (Fig. 16K). Mouth hook: distal blade shorter than basal portion, apically more or less roundish, gently curved downward, with two lines of blunt teeth on submedial portion of ventral margin (Fig. 16L). Holotype. ♂ (MZB), “ Soppeng, South Sulawesi, Indonesia, 9.i.2005, K. T. Takano / Emerged from puparium collected from an inflorescence of Alocasia balgooyi ”. Paratypes. Indonesia: 3♂, 4♀, same data as the holotype (MZB, SEHU). Distribution. Sulawesi (South Sulawesi). Remarks. This species is peculiar in having many unusual characters as described in the diagnosis. Especially, the absence of any process on the male abdominal sternite VI is exceptional in the C. cristata species group. Etymology. Referring to the lack of process on the male abdominal sternite VI.

2.3 Colocasiomyia sabahana species complex, new Diagnosis. Cibarial, posterior sensilla absent (ch.6-1; Fig. 4C). Phallal sheath gently curved dorsad and slightly projected triangularly at dorsobasal corner in lateral view (ch.40-2, 41-1; Figs 11F, 24I). Shared characters. Supracervical setae approximately 4 per side, distally more or less curved, longer than inner occipital setae. Anterior reclinate orbital seta small, situated behind proclinate orbital seta; additional, interfrontal setulae on fronto-orbital plate present (ch.2-0). Eye with stout, dense interfacetal setulae (ch.3-1). Distance between antennal sockets wider than half of socket width (ch.4-0). Cibarial, medial sensilla 3–4 per side, in parallel rows narrower than sensilla campaniformia (ch.5-2; Fig. 4C). Anterior dorsocentral setae behind transverse suture (ch.7- 0). Prescutellar acrostichal setae absent (ch.8-1); acrostichal setulae in 4 rows (ch.9-0). Katepisternal setae small, shorter than postpronotal setae (ch.10-1). Apical scutellar setae nearer to each other than to basal scutellar seta. Costal setae in middle row all weak, trichoid (ch.12-2). Patch covered with only minute pubescence absent on anterodorsal portion of fore tarsomere I (ch.13-1). Male abdominal sternites III–V wider than long (ch.16,17,18-0); V medially concaved on posterior margin. Female abdominal sternite III as long as wide (ch.19-0); V longer than wide (ch.21-1). Male abdominal sternite VI not pubescent at least partly, with a pair of divergent processes longer than 0.1 mm, widely separated from each other, bearing prominent seta on apex and small setae on submedial to distal portion (ch.23,25-0, 24,26,27-1; Fig. 9L,M). Epandrium pubescent except for anterior to ventral margin and anteroventral elongation; setae on posteroventral portion as thick as cercal setae (ch.32-0); phragma small (narrower than epandrial anteroventral corner) expansion on antero-subapical margin (ch.33-2). Surstylus vestigial, very narrow process, apically with 1 or 2 tiny, trichoid seta(e) (ch.34,35-1). Cercus pubescent only on dorsal half; ventral/apical portion more or less differentiated from upper portion, with many small setae (ch.38-0). Phallal sheath not pubescent, as thick as wide, apically not bilobed (ch.42-0, 43,44-1; Figs 11F,G, 24I,J); phallapodeme sclerotized plate directed nearly along or slightly oblique to phallal axis (ch.45-1, 46-0; Figs 11F, 24I). Gonocoxites long, narrow slips (Figs 11F, 24I). Epiproct and hypoproct with pubescence (ch.48-0). Oviscapt shorter than phallus (apodeme + sheath), with ovisensilla only on distal portion occupying less than 1/2 of total length (ch.49-0, 51-1). Puparium (3rd instar larva): segments with stout spicules on ventral surface; anterior spiracle with stalk ending in a whorl of 19–25 long branches; stalk as long as posterior spiracle and longer than longest branch; caudal segments not so elongate, with many small spicules, ending in a V-shaped pair of long posterior spiracles (Fig. 24M,N). Mouth hook moderately, triangularly expanded medioventrally in lateral view; distal blade as long as basal portion, apically pointed, gently curved downward, with two rows of small, acute teeth on submedial to subapical portion of ventral margin (Fig. 24O,P). Included species. Colocasiomyia sabahana sp. nov. and C. sarawakana sp. nov., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on page 37, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402

14) Colocasiomyia sumatrana Toda & Takano, sp. nov. (Figs 9Q, 15M,N, 28) Colocasiomyia sp.5 aff. diconica: Sultana et al., 2006: 694. Diagnosis. All coxae and femora entirely yellow (Fig. 15M,N). A pair of processes on male abdominal sternite VI longer than 0.1 mm, proximally parallel, distally divergent (Fig. 9Q). Description (♂ and ♀; not repeating characters common to C. xenalocasiae). Head. Supracervical setae 9–11 per side. Eye with fine, sparse interfacetal setulae. Cibarial, medial sensilla 2–3 per side; posterior sensillum 1 per side. Supralateral seta outside prementum 1 per side. Thorax. Prescutellar acrostichal setae absent. Apical scutellar setae nearer to each other than to basal scutellar seta. Abdomen. Male tergites: I and II medially pale grayish yellow, laterally dark grayish brown; III–VI nearly entirely dark grayish brown (Fig. 15M). Female tergites variable in color: nearly entirely pale yellow or each tergite with grayish brown to dark gray bands on sublateral portions (Fig. 15N). Male abdominal sternites III and IV longer than wide, rectangular (Fig. 28A). Female abdominal sternites VI longer than wide (Fig. 28B). Male terminalia. Epandrium pubescent except for anterior and ventral margins and anteroventral elongation, with approximately 3 setae on lateral to dorsal portion and 16–17 setae on ventral portion of each side (Fig. 28C). Surstylus with 1–3 recurved, thick, trichoid setae apically and 1–2 minute setae subapically (Fig. 28C). Cercus pubescent except for anterior margin and ventral 2/5, with approximately 25–27 setae (Fig. 28C). Female terminalia. Oviscapt about 5 times as long as wide, slightly curved downward, with 19–20 ovisensilla (Fig. 28F). Indices (range of 10♂ and 10♀ paratypes): FW/HW = 0.55–0.61, ch/o = 0.42–0.58, prorb = 0.59–0.83, rcorb = 0.13–0.27, vb = 0.31–0.53, dcl = 0.52–0.67, sctl = 0.59–0.77, sterno = 0.40–0.63, orbito = 0.43–0.70, dcp = 0.90–1.13, sctlp = 0.90–1.24, C = 1.97–2.59, 4c = 0.90–1.25, 4v = 1.72–2.03, 5x = 1.27–1.73, ac = 2.34–3.60, M = 0.49–0.65, C3F = 0.09–0.29. Holotype. ♂ (MZB), “ Batang Anai, Padang Panjang, West Sumatra, Indonesia, 150 m a.s.l., 5.i.2004, ex Colocasia esculenta, K.T. Takano ”. Paratypes. Indonesia: 14♂, 14♀, same data as the holotype (MZB, SEHU). Distribution. Sumatra (West Sumatra). Remarks. In the cladogram (Fig. 13), this species and the foregoing species C. diconica formed a clade (BP = 75) supported by two synapomorphies: eye with fine, sparse interfacetal setulae (ch.3-0) and male abdominal sternite VI with a pair of processes proximally parallel but distally divergent (ch.29-1). Etymology. Referring to the type locality., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 44-45, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x"]}

13) Colocasiomyia diconica (Toda & Okada, 1983) (Figs 5D, 9P, 15L, 27) Drosophilella diconica Toda & Okada, 1983: 172. Colocasiomyia diconica: Okada, 1988: 36. Diagnosis. Acrostichal setulae in 2 rows (Fig. 5D). A pair of processes on male abdominal sternite VI shorter than 0.1 mm, proximally parallel, distally divergent (Fig. 9P). Supplementary description (not repeating characters common to C. xenalocasiae). Supracervical setae 3–5 per side. Eye with fine, sparse interfacetal setulae. Cibarial, medial sensilla 2–3 per side; posterior sensilla 1–2 per side. Supralateral seta outside prementum 1 per side. Prescutellar acrostichal setae absent (Fig. 5D). Costal setae in middle row all weak, trichoid. Mid tibia with approximately 2 apical, stout setae. Male abdominal sternite III longer than wide, rectangular; IV as wide as long (Fig. 27A). Epandrium pubescent except for anterior margin and anteroventral elongation, posteroventrally less extended, with 13–14 setae on ventral portion of each side (Fig. 27C). Surstylus long, distally tapering, apically truncate plate with 3 recurved teeth apically and 1 minute seta subapically (Fig. 27C). Cercus pubescent except for anterior margin and ventral 1/4 (Fig. 27C). Oviscapt about 4 times as long as wide, with 11–14 ovisensilla (Fig. 27F). Specimens examined. Thailand: 1♂, 2♀, near Chao Phraya River, Taykoa, Bangkok, 14°07’25”N 100°31’18”E, 16.i.2020, ex Colocasia esculenta, P.J. Matthews leg. (SEHU). Malaysia: 28♂, 37♀, Ulu Gombak, Selangor, 8.xii.2013, ex C. esculenta, M.J. Toda leg. (SEHU, UMKL); 12♂, 10♀, Sampadi, Sarawak, 9.viii.2012, ex C. esculenta, A.M. Paulus & K.T. Takano leg. (RDID). Distribution. Borneo (Sabah, Sarawak *, West Kalimantan), Sulawesi?, Java, Sumatra, Peninsular Malaysia *, Vietnam, Thailand, Myanmar, Sri Lanka?., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 42-43, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Toda, M. J. & Okada, T. (1983) Ecological studies of floricolous Drosophilella in Burma with descriptions of three new species from Burma and the Philippines (Diptera, Drosophilidae). Kontyu, 51, 169 - 184.","Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34 - 39."]}

3) Colocasiomyia vieti Toda, sp. nov. (Figs 9F, 15C, 18) Colocasiomyia sp.2 aff. diconica: Sultana et al., 2006: 694. Diagnosis. Male abdominal sternite VI with a pair of long processes nearly straight and divergent (Fig. 9F). Description (♂ and ♀; not repeating characters common to C. grandis sp. nov.). Head. Supracervical setae 4–6 per side. Thorax. Anterior dorsocentral setae behind transverse suture. Prescutellar acrostichal setae absent; acrostichal setulae in 2 or 4 rows. Apical scutellar setae slightly more distant to each other than to basal scutellar seta. Legs. Patch covered with only minute pubescence absent on anterodorsal portion of fore tarsomere I. Abdomen. Male: sternites III and IV wider than long. Female: sternites III–V longer than wide; VI as wide as long, posteriorly dilated, medially notched on posterior margin. Male terminalia. Epandrium pubescent except for narrow anterior to ventral margin and anteroventral elongation, roundish on posteroventral margin (Fig. 18A). Surstylus long, moderately broad plate with 5–6 recurved teeth apically and 1 trichoid seta ventrosubapically (Fig. 18A). Median piece of subepandrial sclerite medially and anteriorly bilobed into long lobes. Cercus pubescent except for anterior margin and apical portion, slightly constricted ventrosubapically, with 34–36 setae all over and small setae on apicoventral margin (Fig. 18A). Phallal sheath apically broadly round in lateral view (Fig. 18B). Female terminalia. Oviscapt with 16–18 ovisensilla (Fig. 18D). Indices (range of holotype and 3♂ and 3♀ paratypes): FW/HW = 0.54–0.57, ch/o = 0.43–0.56, prorb = 0.86– 1.03, rcorb = 0.44–0.52, vb = 0.34–0.47, dcl = 0.50–0.73, sctl = 0.65–0.83, sterno = 0.58–0.78, orbito = 0.64–0.89, dcp = 0.63–0.94, sctlp = 0.62–0.95, C = 2.08–2.39, 4c = 0.89–1.04, 4v = 1.46–1.70, 5x = 0.90–1.13, ac = 2.81–3.14, M = 0.36–0.47. Puparium (3rd instar larva). Anterior spiracle with stalk ending in a whorl of 9–11 branches; stalk longer than longest branch; caudal segments elongate, ending in Y-shaped posterior spiracles (Fig. 18E–G). Mouth hook: distal blade as long as basal portion, apically pointed, gently curved downward, with small, acute teeth (Fig. 18H). Holotype. ♂ (SEHU), “ Cuc Phuong, Nho Quan District, Vietnam, 2.iv.2000, ex Alocasia atropurpurea, M. Yafuso ”. Paratypes. Vietnam: 3♂, 3♀, same data as the holotype (SEHU). Distribution. Vietnam. Remarks. This species resembles C. grandis sp. nov. in many characters, but can be distinguished from it by the diagnostic character and the absence of patch covered with only minute pubescence on anterodorsal portion of fore tarsomere I. Etymology. Patronym, dedicated to Dr. B. T. Viet who helped MY with field work in Vietnam.

15) Colocasiomyia leucocasiae Toda, sp. nov. (Figs 9R, 15O, 29) Colocasiomyia sp.1 aff. diconica: Sultana et al., 2006: 694; Takano et al., 2011: 22; Fartyal et al., 2013, Fig. 6. Diagnosis. Apically blunt, heavy, peg-like costal setae in middle row interspersed with weak, trichoid ones (Fig. 29A). A pair of processes on male abdominal sternite VI nearly straight and parallel (Fig. 9R). Surstylus as broad as ventro-apical lobe of cercus, with 5–6 recurved teeth apically and 1–2 minute setae subapically (Fig. 29D). Epiproct and hypoproct large; hypoproct thicker than width of oviscapt in lateral view (Fig. 29G). Description (♂ and ♀; not repeating characters common to C. xenalocasiae). Head. Supracervical setae 5–6 per side. Cibarial, posterior sensillum 1 per side. Supralateral seta outside prementum 1 per side. Legs. Mid tibia with approximately 2 apical, stout setae. Abdomen. Male abdominal sternites III and IV wider than long, medially notched on posterior margin (Fig. 29B). Female abdominal sternite VI wider than long, medially notched on posterior margin (Fig. 29C). Male terminalia. Epandrium pubescent except for anterior and ventral margins and anteroventral elongation, posteroventrally less extended, with 1–3 setae on lateral to dorsal portion and 16–19 setae on ventral portion of each side (Fig. 29D). Cercus pubescent except for anterior margin and ventral 1/5, with 37–41 setae (Fig. 29D). Oviscapt about 4 times as long as wide, with 12–13 ovisensilla (Fig. 29G). Indices (range of 10♂ and 10♀ paratypes): FW/HW = 0.50–0.71, ch/o = 0.31–0.67, prorb = 0.93–1.16, rcorb = 0.18–0.32, vb = 0.27–0.54, dcl = 0.46–0.70, sctl = 0.63–0.83, sterno = 0.34–0.63, orbito = 0.37–0.74, dcp = 0.79–1.23, sctlp = 0.80–1.02, C = 1.71–2.46, 4c = 1.14–1.53, 4v = 1.79–2.47, 5x = 1.31–2.13, ac = 2.68–3.51, M = 0.40–0.74. Puparium (3rd instar larva). Segments with stout spicules on ventral surface; anterior spiracle sessile, with a bundle of short branches; caudal segments elongate, with small spicules, ending in a pair of posterior spiracles (Fig. 29H–J). Mouth hook triangularly expanded medioventrally in lateral view; distal blade shorter than basal portion, apically more or less roundish, strongly curved downward, with two lines of blunt teeth on submedial portion of ventral margin (Fig. 29K). Holotype. ♂ (MZB), “ Bogor Botanical Garden, Bogor, West Java, Indonesia, 9.i.2004, ex Leucocasia gigantea, K.T. Takano ”. Paratypes. Indonesia: 11♂, 11♀, same data as the holotype; 10♂, 10♀, Madakaripura waterfall, Probolinggo, East Java, 29.xii.2003, ex L. gigantea, K.T.Takano leg.; 10♂, 10♀, Kerinci National Park, Jambi, Sumatra, 7.xii.2004, ex L. gigantea, K.T. Takano leg. (MZB, SEHU). Malaysia: 1♂, Gua Ikan, Dabong, Kuala Krai, Kelantan, 5°21'15"N 102°01'37"E, 15.iii.2016, ex L. gigantea, S.Y. Wong leg.; 10♂, 10♀, ditto except 17.iii.2016 (SEHU). Another specimen examined. Indonesia: 1♂, Sungai Lubuk Paraku, West Sumatra, 970 m a.s.l., 4.i.2004, ex Colocasia esculenta, K.T. Takano leg. (SEHU). Distribution. Peninsular Malaysia, Sumatra (West Sumatra), Java (West and East Java). Remarks. This species morphologically resembles C. diconica in having the epandrium less extended posteroventrally (Figs 27C, 29D) and the short oviscapt about 4 times as long wide (Figs 27F, 29G), but can be distinguished from it by the diagnostic characters. Etymology. Referring to the main host plant. Biogeography, flower-breeding ecology and evolution Based on up-to-date collection records of Colocasiomyia species, the biogeography of the cristata group is currently summarized as follows. Of the six Colocasiomyia species groups, the cristata group is most widely distributed in the Oriental and Papuan regions, covering nearly the entire range of the genus Colocasiomyia from Solomon Islands / Papua New Guinea (east) to Sri Lanka (west) and from Java (south) to Ryukyu Islands/ Taiwan /South China (north) (Fig. 14). The cristata and colocasiae subgroups coexist almost across this range. The three species complexes of the cristata subgroup are more or less different in their distribution ranges. The sabahana complex is endemic to Borneo, and the cristata complex is distributed in the Greater Sunda Islands and Philippines. The xenalocasiae complex is most widely distributed from tropical to subtropical areas of the whole Colocasiomyia range; C. xenalocasiae is confined to the subtropical area, where it shares the same host plant with C. alocasiae of the colocasiae subgroup (Honda-Yafuso 1983; Yafuso, 1994; Toda unpublished data). Of the three species unassigned to any species complex, C. ecornuta and C. vieti are endemic to Sulawesi and Vietnam, respectively, and C. grandis cohabits in the same host-plant inflorescences with C. xenalocasiae and C. alocasiae in northern Vietnam and southern Yunnan (Sultana et al. 2006). Collection records of all species of the cristata group from their host plants at various localities in the Oriental and Papuan regions are compiled in Appendix 8. Modes of host-plant use vary from monopolization by a single species to sharing by 12 species. In ecological observations of this study, we focused on four species of the cristata subgroup, C. cristata and C. sulawesiana of the cristata complex, C. leucocasiae of the xenalocasiae complex and C. sabahana of the sabahana complex, each of which monopolizes its specific host plant. Colocasiomyia cristata was observed using inflorescences/infructescences of Alocasia alba in West Java. When the anthesis started with spathe opening and strong odor emission, adult flies came to the inflorescence at the female flowering phase (Stage II) and stayed in the spathal chamber around the pistillate and the lower intermediate regions of spadix, probably feeding, ovipositing and/or mating there (Fig. 30A). Then, when the spathe began to constrict at the middle portion of intermediate region and pollen release started in the staminate region, the flies crawled up from the spathal chamber to the open staminate region of spadix (Fig. 30B). Eggs were laid mostly on the pistillate region, especially its lower 1/3 section, sporadically on the lower 1/2 intermediate section, and very rarely on the lower 1/3 staminate section (Figs 30C, 31A). Larvae fed in these spadix portions of infructescences at Stages IV and V, and pupariated within the infructescences of Stage V (Fig. 31A). Offspring adults emerged and left the infructescences through a hole at the top of spathal wall (Fig. 30D) before the spathal dehiscence. Colocasiomyia sulawesiana monopolized inflorescences/infructescences of Alocasia macrorrhizos in South Sulawesi. Eggs were laid on the pistillate region and the lower 1/2 intermediate section (Fig. 31B). Young (1st and 2nd instar) larvae were found feeding there and rarely on the lower 1/3 staminate section of spadix in inflorescences/ infructescences at Stages II to IV (Fig. 31B). Older (2nd and 3rd instar) larvae were found exclusively from the pistillate region, most abundantly from its middle 1/3 section, in infructescences of Stage V and pupariated there (Fig. 31B). Adult flies of C. leucocasiae sp. nov. were collected, often together with those of C. gigantea, from inflorescences of Leucocasia gigantea at the Bogor Botanical Garden in West Java. However, Fartyal et al. (2013) revealed that newly eclosed, young C. gigantea flies visit inflorescences of this host plant for feeding, and that the mature flies use inflorescences of Epipremnum pinnatum (L.) Engler (Monsteroideae, Araceae) for breeding. Thus, C. leucocasiae sp. nov. monopolizes inflorescences/infructescences of Leucocasia gigantea for breeding at the Bogor Botanical Garden. Eggs were laid mainly on the pistillate region, most abundantly on its middle 1/3 section, but rarely on the lower 1/2 intermediate section (Fig. 31C). Larvae were confined to the pistillate region, mostly its lower 1/3 section, and pupariated on the lower pistillate region of infructescences at Stage IV (Fig. 31C). Colocasiomyia sabahana sp. nov. monopolized inflorescences/infructescences of Alocasia scabriuscula in the headquarters area of Kinabalu Park, Sabah. Eggs were laid on the pistillate and intermediate regions, more on the latter region, and young (1st and 2nd instar) larvae were found feeding there, more in the former region of inflorescences at Stage IV (Fig. 31D). At Stage V, the dried appendix falls off but the decayed staminate region remains within the spathal capsule of infructescence. Third-instar larvae were observed feeding on the pistillate and staminate regions of some infructescences at Stage V, and empty puparia were found from other Stage-V infructescences, though the numbers of individuals of these pre-imaginal stages were not counted. Thus, all these four species of the cristata subgroup, each of which monopolizes its own host plant for breeding, are pistilicolous, i.e. ovipositing, feeding (larvae) and pupariating mostly on the pistillate (lower, female-flowers) region of spadix, as well as three species (C. pistilicola, C. diconica and C. xenalocasiae) of the xenalocasiae complex, each of which shares its host plant with another stamenicolous species (C. stameniicola, C. colocasiae and C. alocasiae, respectively) of the colocasiae subgroup (Carson & Okada 1980; Toda & Okada 1983; Honda-Yafuso 1983; Yafuso, 1994). In addition, two cristata- subgroup species, C. kotana sp. nov. and C. kinabaluana sp. nov., which share Alocasia macrorrhizos as their common host plant, are both pistilicolous as well, with slight breedingniche differentiation (Takano et al. 2012). Based on the host-use records compiled in Appendix 8, the cristata group is regarded as specializing on the following four genera of the subfamily Aroideae: Alocasia (11 spp.), Colocasia (7 spp.), Leucocasia (1 sp.) and Steudnera (1 sp.). To estimate evolutionary changes and/or acquirement of host plants at the generic level in the cristata group, the four host-plant genera were mapped on the phylogenetic tree having resulted from the morphological grafting cladistic analysis. ACCTRAN and DELTRAN resulted in the same inference of ancestral states (Fig. 32). All the species of the gigantea group (outgroup) use exclusively monsteroid plants for breeding (Fartyal et al. 2013; Li et al. 2014; Jiao et al. 2020). The most recent common ancestor (MRCA) of the cristata group was estimated to have acquired Alocasia as its host plant. Nine extant species of the cristata subgroup use solely Alocasia for breeding with pistilicolous habits so far as known (five of the nine species). Therefore, the MRCA of the cristata group would have been pistilicolous as well. Of the nine Alocasia specialists, eight species are monoxenous, each depending on a single host-plant species; Alocasia macrorrhizos is the most important host plant, being used by four of them (Appendix 8). But, C. sarawakana sp. nov. is polyxenous, having been recorded from four Alocasia species (Appendix 8). After the acquirement of Alocasia as host plant by the MRCA of the cristata group, host change from Alocasia to Colocasia was estimated to have occurred on two branches, i.e. in the MRCAs of the colocasiae subgroup and of the xenalocasiae complex; all the descendant, extant species of these ancestors are known to visit Colocasia inflorescences. Three pairs each consisting of one stamenicolous colocasiae- subgroup species and one pistilicolous xenalocasiae- complex species cohabit in the same host inflorescence: C. stamenicola and C. pistilicola (Carson & Okada 1980), C. colocasiae and C. diconica (Toda & Okada 1983), and C. alocasiae and C. xenalocasiae (Honda- Yafuso 1983; Yafuso, 1994). Okada (1980) hypothesized that this cohabitation was established by a pair of ancestors of the colocasiae subgroup and of the xenalocasiae complex, and that such a pair dispersed and differentiated into the three extant pairs in different geographic areas. The cohabitation would have been achieved by micro-allopatric breeding-site differentiation along the spadix of host plant between two species of the ancestral pair: the ancestor of the colocasiae subgroup would have changed its breeding habits from pistilicolous ones to stamenicolous ones, while the ancestor of the xenalocasiae complex would have retained pistilicolous habits. This change of breeding habits in the former may have been forced by the host-plant change or through interspecific competition with the cohabitant species. In this respect, C. kotana sp. nov. and C. kinabaluana sp. nov. of the cristata complex are interesting: they are both pistilicolous and cohabit in the same inflorescence/infructescence of the host plant Alocasia macrorrhizos (Takano et al. 2012). However, C. kinabaluana sp. nov. tends to be more stamenicolous than C. kotana sp. nov., having the following traits: the longer oviscapt (Figs 21F, 22F), a larger aggregation of adult flies in the upper part of spathal chamber and more eggs deposited on the intermediate region. This situation may or may not correspond to the initial phase of cohabitation of the colocasiae -subgroup and xenalocasiae -complex ancestors. Another possible case of host-plant change is of C. leucocasiae sp. nov. Although only one male specimen of this species has been collected, together with C. diconica, C. sumatrana sp. nov., C. colocasiae and C. iskandari, from an inflorescence of Colocasia esculenta in West Sumatra, this species is very abundantly collected from inflorescences of Leucocasia gigantea (Fig. 31C) and monopolizes it for breeding. Thus, this species may have changed its host plant from Colocasia to Leucocasia. Acquirement of additional host-plant genera, i.e. niche expansion, was estimated to have occurred in four species (Fig. 32). Of the three host-sharing pairs, the pair of C. alocasiae and C. xenalocasiae expanded their host plants to Alocasia and Leucocasia in their northernmost distribution area (subtropical zone), probably due to the scarceness of Colocasia inflorescences there. In addition, C. grandis sp. nov. joined this pair and formed a particular system of three-species cohabitation in a small area restricted to northern Vietnam and southern Yunnan. Within the colocasiae subgroup, which principally depends on Colocasia, C. steudnerae has acquired Steudnera colocasiifolia as another host plant and monopolistically uses its inflorescences/infructescences for breeding (Takenaka et al. 2006). Interestingly, probably in connection with the monopolization of this new host plant, C. steudnerae exhibits a mixture of pistilicolous and stamenicolous breeding habits: ovipositing exclusively on the pistillate region (pistilicolous habit), larvae feeding on decayed tissues of the staminate region (stamenicolous) and pupariating out of the infructescence (stamenicolous). Furthermore, a morphological adaptation for the first pistilicolous habit is seen in the oviscapt: C. steudnerae has the distally broad oviscapt, as well as other pistilicolous species of the cristata subgroup, which seems to be adaptive for laying eggs between pistils placed more loosely. In contrast, other stamenicolous species of the colocasiae subgroup have the distally very narrow oviscapts to insert their eggs into narrow space between stamens packed compactly. With regard to the evolution of these mixed habits and morphology, Takenaka et al. (2006) proposed two hypotheses. The first is that C. steudnerae has secondarily evolved the pistilicolous oviposition habit and the related morphology of oviscapt under the condition lacking a pistilicolous cohabitant species on the new host plant Steudnera colocasiifolia. The second hypothesis is that the mixed situation in C. steudnerae represents an initial, transitional phase of evolution from ancestral, pistilicolous habits to stamenicolous ones in the colocasiae subgroup. The results of phylogenetic analyses in the present study lend support to the first hypothesis. However, more evidence is needed to fully depict the evolution of breeding habits in the colocasiae subgroup: a total of 14 undescribed species of this subgroup, most of which are sympatric with C. steudnerae in northern Vietnam and southern Yunnan (Fig. 14), are awaiting taxonomical, phylogenetical and ecological studies on them. Moreover, cohabiting mechanisms in some systems where more than two Colocasiomyia species are incorporated have remained unexplored at all.

10) Colocasiomyia sarawakana Toda & Yafuso, sp. nov. (Figs 4C, 9M, 15J, 24A,B,D,F,H–J,L,N,P,R) Colocasiomyia sp.3 aff. diconica: Sultana et al., 2006: 694; Toda & Lakim, 2011: 263; Takano et al., 2011: 26. Diagnosis. Additional dorsocentral setae on presutural area present (Fig. 24B).A pair of processes on male abdominal sternite VI nearly straight (Fig. 9M). Epandrium roundish on posteroventral margin; anteroventral elongation broader than cercus in lateral view (Fig. 24H). Female abdominal sternite VI longer than wide (Fig. 24F). Oviscapt narrower than hypoproct, apically somewhat pointed in lateral view (Fig. 24L). Description (♂ and ♀). Head. Supralateral setae outside prementum 3–4 per side. Thorax. Basal scutellar setae long; posterior tip beyond half of apical scutellar setae. Legs. Mid tibia with 2–3 apical, stout setae. Abdomen. Female sternite IV wider than long (Fig. 24F). Male terminalia. Epandrium with 9–10 setae on posteroventral portion (Fig. 24H). Median piece of subepandrial sclerite broad, somewhat quadrate plate; lateral pieces absent. Cercus with 37–38 setae, extended below, apically forming somewhat broad lobe with many small setae (Fig. 24H). Female terminalia. Oviscapt with approximately 12 ovisensilla (Fig. 24L). Indices (range of 10♂ and 10♀): FW/HW = 0.57–0.63, ch/o = 0.31–0.53, prorb = 0.81–1.07, rcorb = 0.20–0.60, vb = 0.20–0.36, dcl = 0.54–0.74, sctl = 0.63–0.89, sterno = 0.43–0.80, orbito = 0.36–0.69, dcp = 0.78–1.25, sctlp = 1.10–1.67, C = 2.12–2.53, 4c = 0.88–1.01, 4v = 1.44–2.11, 5x = 0.81–1.25, ac = 2.06–2.62, M = 0.31–0.50. Holotype. ♂ (RDID), “ Kuching, Sarawak, Malaysia, 27.xi.2004, ex Alocasia macrorrhizos, Lucy Chong & K.T. Takano ”. Paratypes. Malaysia: 11♂, 12♀, same data as the holotype (RDID); 3♂, 3♀, Poring, Mt. Kinabalu, Sabah, 6°02’46”N 116°42’09”E, 29.vii.2004, ex Alocasia scabriscula, K.T. Takano leg. (BORN, KPSP, SEHU). Distribution. Borneo (Sarawak, Sabah, West Kalimantan). Etymology. Referring to the type locality., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on page 39, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Sultana, F., Hu, Y. G., Toda, M. J., Takenaka, K. & Yafuso, M. (2006) Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology, 31, 684 - 702. https: // doi. org / 10.1111 / j. 1365 - 3113.2006.00344. x","Toda, M. J. & Lakim, M. B. (2011) Genus Colocasiomyia (Drosophilidae: Diptera) in Sabah, Bornean Malaysia: high species diversity and use of host aroid inflorescences. Entomological Science, 14, 262 - 270. https: // doi. org / 10.1111 / j. 1479 - 8298.2011.00452. x","Takano, T. K., Suwito, A., Gao, J. J. & Yin, J. T. (2011) Molecular phylogeny of the cristata species group of the genus Colocasiomyia (Diptera: Drosophilidae). Low Temperature Science, 69, 19 - 28."]}

12) Colocasiomyia xenalocasiae (Okada, 1980) (Figs 4A, 6A, 9O, 10F–H, 11D,E, 15K, 26) Drosophilella xenalocasiae Okada, 1980: 218. Colocasiomyia xenalocasiae: Okada, 1988: 36. Drosophilella colocasiae: Okada, 1975: 356 (nec Duda, 1924). Diagnosis. A pair of processes on male abdominal sternite VI convergent (Fig. 9O). Supplementary description. Supracervical setae 5–10 per side. Eye with stout, dense interfacetal setulae. Cibarial, medial sensilla approximately 3 per side; posterior sensillum 0–1 per side (Fig. 4A). Supralateral seta outside prementum 0–1 per side (Fig. 26A). Prescutellar acrostichal setae present; acrostichal setulae in 4 rows. Apical scutellar setae nearly equidistant from each other and from basal scutellar seta. Costal setae in middle row all apically blunt, heavy, peg-like (Fig. 6A). Mid tibia with approximately 3 apical, stout setae. Male abdominal sternites III and IV longer than wide, posteriorly slightly narrowing (Fig. 26B). Female abdominal sternites VI as wide as long, posteriorly dilated, medially notched on posterior margin (Fig. 26C). Epandrium pubescent except for anterior margin and anteroventral elongation, posteroventrally prominently extended, with approximately 1 seta on lateral portion and 11–12 setae on posteroventral portion of each side (Fig. 10F). Surstylus long, narrow plate with 3 recurved teeth apically and 1 minute seta subapically (Fig. 10H). Median piece of subepandrial sclerite medially and anteriorly bilobed into long lobes; lateral pieces long, narrow (Fig. 10G). Cercus pubescent except for anterior margin and ventral 1/3, with approximately 30 setae (Fig. 10F). Oviscapt about 8 times as long as wide, with 16–17 ovisensilla (Fig. 26D). Specimens examined. Japan: 10♂, 10♀, Sembaru, Okinawa, Ryukyu, 29.iv.2007, ex Alocasia odora, M.J. Toda leg. (SEHU). China: 9♂, 11♀, Menglun, Xishuangbanna, Yunnan, 940 m a.s.l., 5.xii.2018, ex Colocasia esculenta, S. Ling leg.; 16♂, 24♀, ditto except for 5.xi.2018, ex Leucocasia gigantea (SEHU). Philippines: 3♂, 5♀, B. Santiago, Iriga, Luzon, 29.xi.2012, ex C. esculenta, P.J. Matthews, M. Medecilo & J.R. Castillo leg. (MPMP, SEHU). Distribution. Ryukyu Is., Philippines * (Luzon), China (Taiwan, Guangdong, Guangxi, Yunnan), Vietnam. * New record., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 40-42, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Okada, T. (1980) Synhospitalic evolution of the genus Drosophilella Duda (Diptera, Drosophilidae), with description of a new species from Okinawa and Taiwan. Kontyu, 48, 218 - 225.","Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34 - 39.","Okada, T. (1975) The Oriental drosophilids breeding in flowers. Kontyu, 43, 356 - 363.","Duda, O. (1924) Beitrag zur Systematik der Drosophiliden unter besonderer Berucksichtigung der palaarktischen u. orientalischen Arten (Dipteren). Wiegmann's Archiv fur Naturgeschichte, 90 (A), 3, 172 - 234, 7 pls."]}

5) Colocasiomyia sulawesiana Okada & Yafuso, 1989 (Figs 9I, 15E, 20) Colocasiomyia sulawesiana Okada & Yafuso, 1989: 48. Diagnosis. A pair of processes on male abdominal sternite VI short (Supplementary description (not repeating characters common to C. cristata). Supracervical setae 5–9 per side. Cibarial, medial sensilla approximately 2 per side. Supralateral seta outside prementum approximately 5 per side. Male abdominal sternite III longer than wide; IV posteriorly slightly widening (Fig. 20A). Female abdominal sternite VI as wide as long, medially concaved on posterior margin (Fig. 20B). Epandrium with 1–3 setae on lateral to dorsal portion and 22–23 setae thicker than cercal setae on ventral portion of each side (Fig. 20C). Cercus with approximately 34–39 setae (Fig. 20C). Oviscapt distally slightly curved ventrad, longer than phallus (apodeme + sheath), with 20–22 ovisensilla but no patch of pubescence (Fig. 20F). Puparium (3rd instar larva): segments with stout spicules on ventral surface; anterior spiracle sessile, with a bundle of approximately 4 short branches; caudal segments elongate, with many small spicules, ending in a V-shaped pair of posterior spiracles (Fig. 20G,H). Mouth hook less expanded medioventrally in lateral view; distal blade as long as basal portion, apically pointed, weakly curved downward, with two rows of small, acute teeth on submedial to subapical portion of ventral margin (Fig. 20I). Specimens examined. Indonesia: 50♂, 37♀, Enrekang, South Sulawesi, 6.i.2005, ex Alocasia macrorrhizos (L.) G. Don, K. T. Takano leg.; 26♂, 22♀, ditto except 8.i.2005 (MZB, SEHU). Distribution. Sulawesi (South Sulawesi). Remarks. Some original descriptions by Okada & Yafuso (1989) should be revised as follows. Okada & Yafuso (1989) described, “Second tarsal joint of fore leg protruded, with about 5 black teeth in 2 rows”, but illustrated four teeth in “ Fig. 1 ” and mentioned, “second tarsal joint of fore leg with about 4 teeth”, in the paragraph of ‘ Relationships ’. The latter state should be correct, because all the specimens examined in this study have four pegs on the fore tarsomere II without any exception. The original descriptions of “male 6S laterally divided, without protuberances” and “Surstylus absent” may be due to overlook of these small or vestigial organs in this species., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 30-32, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Okada, T. & Yafuso, M. (1989) The genus Colocasiomyia Duda (Diptera, Drosophilidae) from Sulawesi. Proceedings of the Japanese Society of Systematic Zoology, 39, 48 - 55."]}

4) Colocasiomyia cristata de Meijere, 1914 (Figs 3E–H, 4B, 5C,F, 6C, 7B,D, 9G, 10C–E, 11H,I, 12C,D, 15D, 19) Colocasiomyia cristata de Meijere, 1914: 273; Duda, 1924: l77; Okada, 1988: 35. Diagnosis. A pair of processes on male abdominal sternite VI long (> 0.05 mm), slightly divergent and curved inward (Fig. 9G). Phallal sheath apically looking like arrowhead in ventral view (Fig. 11I). Supplementary description. Supracervical setae 6–9 per side, nearly straight, as long as inner occipital setae (Fig. 3E). Cibarial, medial sensilla 2–3 per side; posterior sensillum 1 per side (Fig. 4B). Supralateral seta outside prementum 4–5 per side (Fig. 19B). Anterior dorsocentral setae just beside transverse suture (Fig. 5C). Male abdominal sternite III as wide as long; IV wider than long (Fig. 19C). Female abdominal sternites IV–VI longer than wide (Fig. 19D). Epandrium pubescent except for anterior margin and anteroventral elongation, roundish on posteroventral margin, with approximately 2 setae on dorsal portion and 23–24 setae thicker than cercal setae on ventral portion of each side (Fig. 10C). Cercus pubescent on dorsal half but not on anterior margin and ventral portion, extended below, slightly constricted ventrosubapically in lateral view, with approximately 42 setae (Fig. 10C). Phallapodeme less sclerotized thin plate, nearly perpendicular to phallal axis (Fig. 11H). Hypoproct not pubescent (Fig. 12C,D). Oviscapt gently sinuate, as long as phallus (apodeme + sheath), apically round, with patch of pubescence on subdistal portion and 20–21 ovisensilla only on distal portion occupying less than 1/2 length of oviscapt (Fig. 12C). Specimens examined. Indonesia: 1♂, Bogor Botanical Garden, Bogor, West Java, 6°36´9.9˝S, 106°47´47.8˝E, 273 m a.s.l., 9.i.2004, ex Alocasia alba Schott, K.T. Takano leg.; 2♀, ditto except 14.i.2004; 1♂, KPH Gadung, Limus Nunggal, Cisaga, Ciamis, West Java, 7°20'13˝S, 108°32'23˝E, 50 m a.s.l., 22.xii.2004, ex Alocasia alba, K.T. Takano leg.; 5♂, 1♀, Curug Sewu, Central Java, 16.x.2005, ex Alocasia alba, M.J. Toda leg. (SEHU). Distribution. Java (West and Central Java)., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 29-30, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["de Meijere, J. C. H. (1914) Studien uber sudostasiatische Dipteren IX. Tijdschrift voor Entomologie, 57, 137 - 275.","Duda, O. (1924) Beitrag zur Systematik der Drosophiliden unter besonderer Berucksichtigung der palaarktischen u. orientalischen Arten (Dipteren). Wiegmann's Archiv fur Naturgeschichte, 90 (A), 3, 172 - 234, 7 pls.","Okada, T. (1988) Taxonomic note on Colocasiomyia cristata de Meijere (Diptera, Drosophilidae) with generic synonymy. Proceedings of the Japanese Society of Systematic Zoology, 37, 34 - 39."]}

8) Colocasiomyia matthewsi Toda, sp. nov. (Figs 9H, 15H, 23) Colocasiomyia sp. aff. cristata: Fartyal et al., 2013: 768. Diagnosis. A pair of processes on male abdominal sternite VI long (> 0.05 mm), widely separated, strongly divergent, slightly curved inward (Fig. 9H). Epandrium pubescent except for anteroventral elongation and anterior, ventral and posteroventral margins, posteroventrally extended as ventral lobe somewhat pointed and bearing many small setae on apex, with 7–8 setae on lateral portion and 30–31 setae as thick as cercal setae on ventral lobe of each side (Fig. 23C). Phallal sheath apicoventrally narrowed and apically round in lateral view (Fig. 23D). Oviscapt very short, only twice as long as wide, somewhat oval, with 23–24 ovisensilla on nearly entire surface (Fig. 23F). Description (♂ and ♀; not repeating characters common to C. kinabaluana sp. nov.). Head. Supracervical setae 5–8 per side, distally more or less curved, longer than inner occipital setae. Cibarial, medial sensilla 3–4 per side. Supralateral setae outside prementum 3–5 per side. Thorax. Anterior dorsocentral setae behind transverse suture. Abdomen. Male sternite III longer than wide, posteriorly slightly narrowing; IV wider than long (Fig. 23A). Female sternite V wider than long; VI wider than long, medially concaved on posterior margin (Fig. 23B). Male terminalia. Cercus pubescent only on dorsal 2/5, with 56–57 setae, extended below, apically forming broad lobe marginally roundish and fringed with small setae (Fig. 23C). Phallapodeme sclerotized plate slightly oblique to phallal axis (Fig. 23D). Female terminalia. Hypoproct medially with a small patch of pubescence (Fig. 23F). Indices (range of 10♂ and 10♀ paratypes): FW/HW = 0.53–0.58, ch/o = 0.28–0.36, prorb = 0.86–1.14, rcorb = 0.33–0.56, vb = 0.26–0.47, dcl = 0.57–0.74, sctl = 0.74–0.98, sterno = 0.65–1.10, orbito = 0.45–0.82, dcp = 0.97–1.30, sctlp = 1.27–1.82, C = 1.74–2.12, 4c = 0.92–1.08, 4v = 1.42–1.60, 5x = 0.72–1.14, ac = 2.60–3.45, M = 0.28–0.40. Holotype. ♂ (MPMP), “ Gandara, Samar, Philippines, 2.xii.2012, ex Alocasia macrorrhizos, P.J. Matthews, M. Medecilo & J.R. Castillo ”. Paratypes. Philippines: 10♂, 10♀, same data as the holotype (MPMP, SEHU). Distribution. Philippines (Samar). Remarks. This species was placed as the basal branch sister to the clade (BP = 100) consisting of the remaining four species of the cristata complex (Fig. 13), lacking the following synapomorphies for the latter clade: anterior dorsocentral setae just beside transverse suture (ch.7-1), epandrium posteroventrally roundish (ch.30-1) and with setae thicker than cercal setae (ch.32-1), phallapodeme less sclerotized thin plate (ch.45-2) directed nearly perpendicular to axis of phallal sheath (ch.46-1), and hypoproct not pubescent (ch.48-1). In addition, within the latter clade, C. sulawesiana, C. kinabaluana sp. nov. and C. kotana sp. nov. formed a clade (BP = 75; Fig. 13) supported by an autapomorphy, the short (C. matthewsi sp. nov. somewhat resembles C. grandis sp. nov. in the morphology of periphallic organs (Figs 16D, 23C), but can be distinguished from it by their diagnostic characters. Etymology. Patronym, dedicated to Prof. Peter J. Matthews, one of the three collectors of the specimens., Published as part of Takano, Kohei Takenaka, Gao, Jian-Jun, Hu, Yao-Guang, Li, Nan-Nan, Yafuso, Masako, Suwito, Awit, Repin, Rimi, Pungga, Runi Anak Sylvester, Meleng, Paulus Ak, Kaliang, Clement Het, Chong, Lucy & Toda, Masanori J., 2021, Phylogeny, taxonomy and flower-breeding ecology of the Colocasiomyia cristata species group (Diptera: Drosophilidae), with descriptions of ten new species, pp. 1-70 in Zootaxa 5079 (1) on pages 34-37, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Fartyal, R. S., Gao, J. J., Toda, M. J., Hu, Y. G., Takano, K. T., Suwito, A., Katoh, T., Takigahira, T. & Yin, J. T. (2013) Colocasiomyia (Diptera: Drosophilidae) revised phylogenetically, with a new species group having peculiar lifecycles on monsteroid (Araceae) host plants. Systematic Entomology, 38, 763 - 782. https: // doi. org / 10.1111 / syen. 12027"]}