Search

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"]}

1. Colocasiomyia colocasiae species subgroup Colocasiomyia colocasiae species subgroup: Takenaka et al., 2006: 84. Diagnosis. Epandrial phragma prominent (wider than epandrial anteroventral corner), somewhat shell-shaped expansion on antero-subapical margin (ch.33-1; Fig. 10A). Surstyli absent (ch.34-2; Fig. 10A). Lateral pieces of subepandrial sclerite less sclerotized, small, somewhat triangular plate-like (ch.37-1; Fig. 10A). Cercus nearly entirely oval (ch.38-1; Fig. 10A). Phallal sheath prominently projected anterodorsad at dorsobasal corner in lateral view (ch.41-2; Fig. 11A). Shared characters. Supracervical setae distally more or less curved, longer than inner occipital setae (ch.1-0; Fig. 3A). Additional, interfrontal setulae on fronto-orbital plate absent (ch.2-1; Fig. 3B). Distance between antennal sockets wider than half of socket width (ch.4-0; Fig. 3C). Cibarial, medial sensilla in parallel rows narrower than sensilla campaniformia (ch.5-2); posterior sensilla present (ch.6-0). Anterior dorsocentral setae situated behind transverse suture (ch.7-0; Fig. 5A). Additional dorsocentral setae on presutural area absent (Fig. 5A). Prescutellar acrostichal setae absent (ch.8-1; Fig. 5A). Acrostichal setulae in 4 rows (ch.9-0; Fig. 5A). Katepisternal setae large; posterior one as long as or longer than longest, postpronotal seta (ch.10-0; Fig. 5E). Basal scutellar setae short; posterior tip not reaching to half of apical scutellar setae (ch.11-0; Fig. 5A). Apical scutellar setae nearly equidistant from each other and from basal scutellar seta (Fig. 5A). Apically blunt, heavy, peg-like costal setae in middle row interspersed with weak, trichoid ones (ch.12-1; Fig. 6B). Fore tarsomere I with patch covered with only minute pubescence on anterodorsal portion (ch.13-0; Fig. 7A). Mid tibia with 2 or more apical setae (ch.15-0; Fig. 7C). Male abdominal sternites IV and V and female sternites III–VI longer than wide (ch.17–22-1; Fig. 8C,D). Male abdominal sternite VI not pubescent at least partly (ch.23-0; Fig. 9B–D), with single process longer than 0.05 mm and not bearing prominent seta on apex but small setae on submedial to distal portion (ch.24-2, 25–27-0; Fig. 9B–D). Epandrium roundish on posteroventral margin (ch.30-1), with seta as long as width of epandrium and short setae as thick as cercal setae on posteroventral portion (ch.31,32-0; Fig. 10A). Median piece of subepandrial sclerite broad, somewhat quadrate plate (ch.36-0; Fig. 10A,B). Cercus wider than 1/2 cercal length (ch.39-1; Fig. 10A). Phallal sheath nearly straight, apically vertically bilobed, as thick as wide, without pubescence (ch.40-0, 42–44-1; Fig. 11A,B). Phallapodeme more or less sclerotized, slightly oblique to axis of phallal sheath, longer than 1/2 of phallal sheath (ch.45-1, ch.46,47-0; Fig. 11A,B). Epiproct and hypoproct pubescent (ch.48-0; Fig. 12A,B). Oviscapt not longer than phallus (apodeme + sheath), with ovisensilla on portion occupying 1/2 or more in length of oviscapt; distal portion differentiated into more or less narrow process from basal, broad portion with dense, fine wrinkles (ch. 49–51-0; Fig. 12A). Included species. Colocasiomyia colocasiae, C. alocasiae, C. iskandari, C. steudnerae and C. stamenicola. A total of 14 putatively new species probably belong to this subgroup (Fartyal et al. 2013; Toda unpublished data; Fig. 14). The species-status determination and description of them will be done elsewhere., 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 20-21, DOI: 10.11646/zootaxa.5079.1.1, http://zenodo.org/record/5766402, {"references":["Takenaka, K., Yin, J. T., Wen, S. Y. & Toda, M. J. (2006) Pollination mutualism between a new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) and Steudnera colocasiifolia (Araceae) in Yunnan, China. Entomological Science, 9, 79 - 91. https: // doi. org / 10.1111 / j. 1479 - 8298.2006.00156. 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"]}

Plant functional traits on serpentine degraded areas in Sabah (Malaysia). A potential tool for its reclamation. International Conferences on Serpentine Ecology (ICSE)

Recovery of soil function in a serpentine quarry in Sabah (Malaysia) along an age gradient of the pioneer tree Ceuthostoma terminale. International Conferences on Serpentine Ecology (ICSE)

Rhizobium nitrogen-fixing nodules are a well-known trait of legumes, but nodules also occur in other plant lineages either with rhizobium or the actinomycete Frankia as microsymbiont. The widely accepted hypothesis is that nodulation evolved independently multiple times, with only a few losses. However, insight in the evolutionary trajectory of nodulation is lacking. We conducted comparative studies using Parasponia (Cannabaceae), the only non-legume able to establish nitrogen fixing nodules with rhizobium. This revealed that Parasponia and legumes utilize a large set of orthologous symbiosis genes. Comparing genomes of Parasponia and its non-nodulating relative Trema did not reveal specific gene duplications that could explain a recent gain of nodulation in Parasponia. Rather, Trema and other non-nodulating species in the order Rosales show evidence of pseudogenization or loss of key symbiosis genes. This demonstrates that these species have lost the potential to nodulate. This finding challenges a long-standing hypothesis on evolution of nitrogen-fixing symbioses, and has profound implications for translational approaches aimed at engineering nitrogen-fixing nodules in crop plants.

Two taxonomically undescribed Colocasiomyia species were discovered from inflorescences of Alocasia macrorrhizos in Kota Kinabalu City, Sabah, Borneo, Malaysia. The aims of this study were to investigate the reproductive ecology of the flies and the plant, ascertain the importance of the flies as pollinators and examine the intimate association between flowering events and life history of the flies. We conducted sampling, observations and field pollination experiments. The flies were attracted by the odour of female-phase inflorescences in the early morning on the first day of anthesis. They fed, mated and oviposited in the inflorescences for one day. On the second day, the flies, covered with pollen grains, left the male-phase inflorescences for the next female-phase inflorescences. The immatures of both fly species hatched, developed and pupated within the infructescences without damaging the fruits and developed adults emerged when the mature infructescences dehisced. The flowering events and fly behaviours were well synchronized. In the field pollination experiments, the inflorescences bagged with a fine mesh (insect exclusion) produced almost no fruits, whereas those bagged with a coarse mesh (bee exclusion) produced as many fruits as the open-pollinated controls. These results indicate that the flies are the most efficient and specialised pollinators for their host, A. macrorrhizos. These flies, in return, depend on A. macrorrhizos for food and habitat through most of their life cycle. This study provides a deeper insight into the less recognised, highly intimate pollination mutualism between Araceae plants and Colocasiomyia flies.