Uncovering hidden diversity: phylogeny and taxonomy of Physoderinae (Reduviidae, Heteroptera), with emphasis on Physoderes Westwood in the Oriental and Australasian regions Author Hwang, Wei Song 1899A130-C5D0-43F1-8299-069E425DC965 Lee Kong Chian Natural History Museum, Department of Biological Sciences, National University of Singapore, 117377, Singapore. & Department of Entomology, University of California, Riverside, Riverside, CA 92521, USA. & urn: lsid: zoobank. org: author: 1899 A 130 - C 5 D 0 - 43 F 1 - 8299 - 069 E 425 DC 965 & Corresponding author: nhmhws @ nus. edu. sg nhmhws@nus.edu.sg Author Weirauch, Christiane 59B6A54F-2044-4E5A-B7F7-AAC5E7D716F0 Department of Entomology, University of California, Riverside, Riverside, CA 92521, USA. & Email: christiane. weirauch @ ucr. edu & urn: lsid: zoobank. org: author: 59 B 6 A 54 F- 2044 - 4 E 5 A-B 7 F 7 - AAC 5 E 7 D 716 F 0 christiane.weirauch@ucr.edu text European Journal of Taxonomy 2017 2017-08-03 341 1 118 journal article 22055 10.5852/ejt.2017.341 5449e1a8-d793-4d90-9b61-1c66964b3af3 2118-9773 3832798 12B0369B-62CC-4D3D-B933-5EF0FA9B49AA Physoderes curculionis China , 1935 Figs 8 , 14 , 16–17 , 19–20 ; Appendix Physoderes curculionis China , 1935: 497 . Physoderes insulanus Miller, 1940: 543 , fig. 88. New synonymy. Physoderes patagiata Miller, 1941: 782 , fig. 7. New synonymy. Physoderes minor Usinger, 1946: 50 , fig. 11. New synonymy. Physoderes kalshoveni Miller, 1954: 86 , fig. 10. New synonymy. Diagnosis This species is recognized among species of Physoderes by its small size ( 7.64 mm average length), the short and conical head, the scape reaching the apex of the clypeus, the anterior pronotal lobe being equal to or slightly narrower than the posterior lobe, the posterior pronotal lobe being rugose, the rounded parascutellar lobe, which is skewed towards the median, the semicircular scutellum, the spatulate and apically rounded scutellar process, the hemelytron not reaching the tip of the abdomen, the anterior margin of mediosternite 8 undulating and with an apodeme, the extremely narrow transverse bridge of the pygophore, and the endosomal struts being apically subacute and basally divided into two plates converging medially to form a ridge. This species is most similar to P. azrael , but can be differentiated by the elevation of the pronotum, the shorter and more ovoid abdomen, and the shape of the endosomal struts. Material examined Holotype MALAYSIA : , Malaya , unknown locality, 3.11° N , 101.72111° E , no date provided, ( UCR_ENT 00018529 ) ( BMNH ). Paratype MALAYSIA : 1 ♀ , Selangor , Petaling Co., Serdang, 3.02277° N , 101.71361° E , 14 Jan. 1935 , G.H. Corbett leg. ( UCR _ENT 00018525) ( BMNH ). Other material CHRISTMAS ISLAND : 1 ♀ , 10.45917° S , 105.68972° E , 248 m , 28 Jan. 1933 , unknown collector, holotype of Physoderes insulanus (junior synonym) ( UCR _ENT 00018532) ( BMNH ). GUAM : 1 ♂ , 3 mi. S of Piti, 13.45444° N , 144.70444° E , 98 m , 23 May 1936 , O.H. Swezey leg., paratype of Physoderes minor (junior synonym) ( UCR _ENT 00073581) ( BPBM ); 1♀ , same collection data as preceding ( UCR _ENT 00073592) ( BPBM ); 1 ♀ , Mariana Is., Mt Lamlam Co., Mt Lalam, 13.33583° N , 144.66527° E , 406 m , Dec. 1958 , N.L.H. Krauss leg. ( UCR _ENT 00073596) ( BPBM ); 1 ♀ , 1 mi. SE of Asan, 13.47305° N , 144.71305° E , 244 m , 31 Oct. 1947 , H.S. Dybas leg. ( UCR _ENT 00031420) ( USNM ); 1 ♀ , Pilgo River, 13.44222° N , 144.77416° E , 26 May 1945 , J.L. Gressitt leg. ( UCR _ENT 00031421) ( USNM ); 1 ♀ , same collection data as preceding but G.E. Bohart and J.L. Gressit leg. ( UCR _ ENT 00073595) ( BPBM ); 2 ♂♂ , Port Ajayan, 13.2725° N , 144.70444° E , 143 m , 6 Jun. 1945 , H.S. Dybas leg. ( UCR _ENT 00073588, UCR _ENT 00073589) ( BPBM ); 1 ♀ , 2 mi. W of Piti, 13.45472° N , 144.695° E , 61 m , on Glycine sp. ( Fabaceae ), 20 Jul. 1937 , R.G. Oakley leg. ( UCR _ENT 00031419) ( USNM ); 3 ♀♀ , Dededo, 13.52° N , 144.84° E , 19 May 1936 , R.L. Usinger leg., paratypes of Physoderes minor (junior synonym) ( UCR _ENT 00046627– UCR _ENT 00046629) ( CAS ); 1 ♀ , Dededo, 13.52° N , 144.84° E , 13 Jul. 1937 , R.G. Oakley leg. ( UCR _ENT 00031416) ( USNM ); 1 ♂ , same collection data as preceding but 23 Feb. 1938 ( UCR _ENT 00031412) ( USNM ); 2 ♀♀ , same collection data as preceding but 25 Aug. 1937 ( UCR _ENT 00031417, UCR _ENT 00031418) ( USNM ); 2 ♀♀ , Harmon Field, 13.50722° N , 144.81527° E , 54 m , on Carica papaya L. ( Caricaceae ), 21 Jan. 1949 , collector unknown ( UCR _ENT 00031422, UCR _ENT 00031423) ( USNM ); 1 ♂ , Mt Alifan, 13.38055° N , 144.67138° E , Jun. 1936 , R.L. Usinger leg., paratype of Physoderes minor (junior synonym) ( UCR _ENT 00073579) ( BPBM ); 1 ♀ , same collection data as preceding but 26 May 1936 , paratype of Physoderes minor (junior synonym) ( UCR _ENT 00068911) ( AMNH ); 1 ♂ , same collection data as preceding, paratype of Physoderes minor (junior synonym) ( UCR _ENT 00046625) ( CAS ); 1 ♀ , Mt Santa Rosa, 13.50805° N , 144.91083° E , 182 m , 16 May 1948 , G.E. Bohart and J.L. Gressit leg. ( UCR _ENT 00031425) ( USNM ); 1 ♂ , Piti, 13.46° N , 144.69166° E , 5 m , 26 May 1936 , O.H. Swezey leg., paratype of Physoderes minor (junior synonym) ( UCR _ENT 00073577) ( BPBM ); 1 ♀ , same collection data as preceding but 27 Oct. 1936 , paratype of Physoderes minor (junior synonym) ( UCR _ENT 00073593) ( BPBM ); same collection data as preceding, paratype of Physoderes minor (junior synonym) ( UCR _ENT 00046626) ( CAS ); 1 juv. , Piti, 13.46° N , 144.69166° E , 5 m , 22 May 1936 , O.H. Swezey leg. ( UCR _ENT 00046624) ( CAS ); 1 ♀ , Pt Oca, 13.503° N , 144.771° E , May 1945 , G.E. Bohart and J.L. Gressit leg. ( UCR _ENT 00073590) ( BPBM ); 1 ♂ , same collection data as preceding but 19 May 1945 ( UCR _ENT 00073587) ( BPBM ); 1 ♀ , same collection data as preceding but 15 May 1945 ( UCR _ENT 00073594) ( BPBM ); 1 ♀ , Santa Rita, 13.37416° N , 144.70805° E , 69 m , 25 Jan. 1948 , collector unknown ( UCR _ENT 00031424) ( USNM ); 1 ♂ , unknown locality, 13.44416° N , 144.79361° E , 37 m , no date provided, D.T. Fullaway leg. ( UCR _ENT 00031413) ( USNM ); 2 ♀♀ , Guam, 1937, R.G. Oakley leg. ( UCR _ENT 00031414, UCR _ENT 00031415) ( USNM ); 2 ♂♂ , Yigo, 13.53194° N , 144.88027° E , 126 m , 13 Nov. 1936 , O.H. Swezey leg., paratypes of Physoderes minor (junior synonym) ( UCR _ENT 00073578, UCR _ENT 00073580) ( BPBM ). Fig. 16. Dorsal habitus images for junior synonym holotypes of species in Physoderes Westwood, 1845 (except P. minor = paratype). Scale bars = 2 mm. INDONESIA : Java : 1 ♂ , Ambarawa, 7.25611° S , 110.40638° E , no date provided, collector unknown ( UCR _ENT 00024014) ( RMNH ); 1 ♂ , Buitenzorg (Bogor), 6.58916° S , 106.79305° E , 250 m , Jan. 1926 , L.G.E. Kalshoven leg., holotype of Physoderes kalshoveni (junior synonym) ( UCR _ENT 00023947) ( RMNH ); 1 ♀ , Bogor, 6.58916° S , 106.79305° E , 240 m , 28 Apr. 1954 , A.H.G. Alston leg. ( UCR _ENT 00069413) ( BMNH ); 1 ♀ , Bogor, 6.5897° S , 106.7914° E , 2 Jul. 1936 , J.v.d. Vecht leg. ( UCR _ENT 00024013) ( RMNH ); 1 ♂ , Samarang, 6.96666° S , 110.41666° E , 4 m , Jun. 1910 , E. Jacobson leg. ( UCR _ENT 00024012) ( RMNH ). – Maluku : 1 ♂ , Ambon County, Waai, Ambon Island, 3.565° S , 128.32083° E , 93 m , 10 Mar. 1965 , A.M.R. Wegner leg. ( UCR _ENT 00073586) ( BPBM ). – Larat: 1 ♀ , 7.78777° S , 131.81694° E , Dec. 1907 , F. Muir leg. ( UCR _ENT 00046621) ( CAS ). – Sumatra : 1 subadult , Sumatera Barat ( West Sumatra ), 0.74° N , 100.8° E , no date provided, Muller leg. ( UCR _ ENT 00024032) ( RMNH ). MALAYSIA : Kuala Lumpur : 1 ♀ , 3.13888° N , 101.68694° E , 43 m , 5 Feb. 1940 , N.C.E. Miller leg., paratype of P. patagiata (junior synonym) ( UCR _ENT 00069396) ( BMNH ). – Penang : 6 ♂♂ , Penang Island, 5.37027° N , 100.2375° E , 198 m , no date provided, Baker leg. ( UCR _ENT 00031397– UCR _ENT 00031402); 8 ♀♀ , same collection data as preceding ( UCR _ENT 00031403– UCR _ENT 00031410) ( USNM ). – Sarawak : 1 ♀ , Kuching, 1.53111° N , 110.34416° E , 12 m , no date provided, collector unknown ( UCR _ENT 00069397) ( BMNH ). NORTHERN MARIANA ISLANDS : 1 ♂ , Saipan , Pagan Island, 18.13555° N , 145.79111° E , 155 m , 22 Aug. 1954 , G. Corwin leg. ( UCR _ENT 00073582) ( BPBM ); 1 ♀ , same collection data as preceding ( UCR _ENT 00073591) ( BPBM ); 1 subadult , same collection data as preceding ( UCR _ENT 00073597) ( BPBM ); 1 juv. , same collection data as preceding ( UCR _ENT 00073598) ( BPBM ). SINGAPORE : 1 ♂ , 1.27166° N , 103.83416° E , 12 m , no date provided, Baker leg. ( UCR _ENT 00031411) ( USNM ). Redescription Male BODY LENGTH. Small, total length 7.64 mm , SD ± 0.21 (Appendix). COLORATION ( Figs 14 , 16–17 ). Brown. Head brown. Scape, pedicel and basiflagellomere of antenna light brown, distiflagellomere basally light brown, straw-colored apically. First segment of labium light brown, second segment straw-colored, third segment brown. Pronotum of thorax anterior lobe light brown to dark brown with straw-colored ridges, posterior lobe light brown to brown, with straw-colored posterior margin including parascutellar lobe, scutellum basally brown, apically straw-colored including scutellar process, pleuron brown to dark brown, sternum brown to dark brown. Corium of hemelytron light brown to brown, membrane dark brown. Fore femora of legs straw-colored, sub-basally and apically with brown annulations, tibiae straw colored, medially and apically with brown annulations, tarsi and claws straw-colored to brown. Abdomen dorsally yellowish-orange, ventrally straw-colored with brown and dark brown patterns, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore brown or dark brown. VESTITURE. Sparsely setose. Head with some curved setae and short, fine, adpressed setae, ventral surface of postocular lobe with sparse, setigerous tubercles, with pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with tuberculated, short, curved setae on lateral margins and along dorsal ridges, posterior lobe with short, curved setae on humeral angle and sparsely distributed along dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibia with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen with a few clubbed setae on each segment, or connexival margin with no prominent setae. HEAD. Short conical; maxillary plate rounded apically; scape reaching apex of clypeus; eye hemispherical in dorsal view, less than 1/5 length of head. THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe with low ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defined; anterior pronotal lobe longer than posterior lobe, equal width to or slightly narrower than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum semicircular, scutellar process long, apex rounded or spatulate. HEMELYTRON. Not reaching tip of abdomen. LEGS. Same as genus description. ABDOMEN. Ovoid, with rounded terminal margin; connexival margin slightly undulating, posterior margin not elevated. GENITALIA. Anterior margin of mediosternite 8 undulating, with medial apodeme; transverse bridge of pygophore extremely narrow, only consisting of margin of anterior opening of pygophore, margin of anterior opening rounded, apodeme absent, apical margin of posterior opening smooth; cup-like sclerite apically rounded with adjacent paired round protuberances; arms of basal plate rounded; ductifer with sclerotized rounded ring; endosomal struts apically subacute, basally divided into two plates converging medially to form a ridge; shape of dorsal phallothecal sclerite subacute, with short, apical lobes and rounded lateral plates at apex. Female Similar to male except slightly narrower anterior pronotal lobe width ( 2.29 mm average) than posterior lobe width ( 2.64 mm average). Ecology This species has been recorded to feed on curculionid larvae and has been collected on decaying plant material including fallen banana ( Musaceae ) logs, rotten papaya ( Caricaceae ) plant, rotten pumpkin ( Cucurbitaceae ), on logs, on rotten boards on the ground and beneath rotten breadfruit ( Moraceae ). Distribution This species has the most widespread distribution of all Oriental physoderines and can be found in peninsular Malaysia , Sumatra, Singapore , Java, Christmas Island , Ambon Island , Guam and Saipan. The presence of P. curculionis in Guam and Saipan is an odd distribution that questions whether this could be a secondary introduction, considering their association with banana, pumpkin and papaya plant material and curculionid larvae. Remarks This species is confirmed to belong to the Physoderes clade based on the phylogenetic analysis above. A number of names have been synonymized under P. curculionis based on examination of the type material. These are P. insulanus , P. kalshoveni , P. patagiata and P. minor . All share the diagnostic characters of P. curculionis listed above including those of the male genitalic morphology (for P. minor and P. kalshoveni ), have no distinct features of their own, and are thus synonymized here.