Uncovering hidden diversity: phylogeny and taxonomy of Physoderinae (Reduviidae, Heteroptera), with emphasis on Physoderes Westwood in the Oriental and Australasian regions

Abstract. The cryptic assassin bug subfamily Physoderinae is characterized by a distinctly uneven species and genus-level diversity across continents, but the lack of a phylogeny has so far precluded investigations into the biogeographic history of the group. Endemic Madagascan Physoderinae (11 of the 15 genera) exhibit broad morphological diversity, but the large (38 spp.) and seemingly uniform genus Physoderes is widely distributed across the Oriental and Australasian regions. The three remaining genera are small or even monotypic and restricted to the Neotropical (Cryptophysoderes, Leptophysoderes) and Afrotropical (Porcelloderes) regions. To investigate relationships among Physoderinae, with emphasis on the monophyly of the Madagascan fauna and the monophyly of Physoderes, we conducted a cladistic analysis based on 57 morphological characters and complete genus-level taxon sampling. We found that the Madagascan fauna is not monophyletic, indicating that the island was colonized more than once, although the great majority of Madagascan taxa are part of a single clade. Overall relationships are recovered as Afrotropical Porcelloderes + [(Neotropical Cryptophysoderes, Leptophysoderes) + (Madagascan, Oriental and Australasian taxa)]. Physoderes is shown to be polyphyletic and 3 new genera, Breviphysoderes gen. nov., Macrophysoderes gen. nov. and Nanophysoderes gen. nov., are erected to accommodate new species and several species previously classified as Physoderes. A taxonomic revision of Physoderes and allied genera is conducted. Diagnoses, updated distribution ranges, habitus and genitalic images, and identification keys are provided. A total of 17 synonymies are given, and 15 new species and 3 new genera are described, focusing on the diversity of Physoderinae in the Oriental and Australasian regions. The new species are Breviphysoderes fulvopicta gen. et sp. nov., B. tenebrosa gen. et sp. nov., Macrophysoderes cirripilosa gen. et sp. nov., M. elongata gen. et sp. nov., M. finisterre gen. et sp. nov., M. grandis gen. et sp. nov., Paraphysoderes popeye sp. nov., Physoderes anamalaiensis sp. nov., P. brevipennis sp. nov., P. minime sp. nov., P. muluensis sp. nov., P. mysorensis sp. nov., P. nigripennis sp. nov., P. ractepilosa sp. nov. and P. tricolor sp. nov.


Introduction
The reduviid subfamily Physoderinae is among the lesser-known groups of assassin bugs due to cryptic habits, restriction of most species to apparently small endemic ranges in wet-tropical areas of the Old and New Worlds, and the overall small number of genera and species compared to other reduviid subfamilies (Rédei 2012;Weirauch et al. 2014): only 70 species classifi ed into 15 genera are described to date (Maldonado 1990;Weirauch 2006;Davranoglou 2014;Davranoglou et al. 2015). All members are small, rarely collected, feature subtle hues of brown and dirty yellow that allow them to blend into leaf litter and decaying wood, and many species share a somewhat dorsoventrally fl attened, compact body shape and relatively short, heavily armored legs (Fig. 1). In contrast, the physoderine fauna of Madagascar and the nearby Comoros Islands (11 genera with 29 spp.;Villiers 1962) shows considerable morphological diversity, with some taxa resembling assassin bugs in other subfamilies, such as Tribelocephaloides Villiers, 1962 andMimoelasmodema Villiers, 1962, that superfi cially resemble Tribelocephalini (in the subfamily Ectrichodiinae) and Elasmodeminae, respectively (Villiers 1962;Weirauch et al. 2014;Forthman & Weirauch 2017;Fig. 2). More speciose, but morphologically relatively uniform, the Oriental and Australasian fauna of Physoderinae comprises 38 species currently classifi ed within a single genus -Physoderes Westwood, 1845 (Maldonado 1990;Cao et al. 2011;Chlond 2011). Only two Neotropical genera of Physoderinae have so far been described, the monotypic Cryptophysoderes Maldonado, 1972 from Panama andLeptophysoderes Weirauch, 2006 from Costa Rica and Ecuador, with L. sarapiqui Davranoglou, Hwang & Weirauch, 2015 showing pronounced sexual dimorphism (Wygodzinsky & Maldonado 1972;Weirauch 2006;Davranoglou et al. 2015). The only described continental Afrotropical species of Physoderinae is Porcelloderes impenetrabilis Rédei, 2012 from mountain ranges in northeastern Tanzania (Rédei 2012).
The present classifi cation of Physoderinae is largely the result of the studies of two researchers: André Villiers and Norman C. E. Miller. Villiers (e.g., 1962 described the majority of the physoderine diversity in Madagascar during the 1950s and 1960s, and Miller (e.g., 1940, 1941 described the Oriental and Australasian Physoderes species in South-East Asia and the Pacifi c from the 1940s to the 1960s. China, 1936 in Singapore on mossy substrate (left) and vegetation debris (right).
Phylogenetic relationships of Physoderinae, both between genera and species within the group and to other clades of assassin bugs, have largely remained in the dark. Miller (1954) suspected that the Neotropical reduviine Aradomorpha Champion, 1899 may be closely related to Physoderinae, whereas Lent & Wygodzinsky (1979) tentatively proposed a relationship with Triatominae, and Weirauch (2008) recovered the African reduviine Sphedanovarus Jeannel, 1866 as the sister taxon of the single species of Physoderes included in her morphology-based analysis. Molecular analyses found a poorly supported relationship with the Neotropical reduviine genus Leogorrus Stål, 1859 as part of an equally poorly supported larger clade also containing other Reduviinae and Salyavatinae (Weirauch & Munro 2009). These hypotheses were either not based on cladistic analyses (Miller 1954;Lent & Wygodzinsky 1979) or hampered by inadequate taxon sampling (Weirauch 2008;Weirauch & Munro 2009). A recent molecular phylogenetic analysis with more comprehensive sampling of Reduviinae (but lacking Aradomorpha) found Physoderinae to be closely related to the Neotropical reduviine genera Microlestria Stål, 1872 and Nalata Stål, 1860(Hwang & Weirauch 2012. To date, relationships among genera and species of Physoderinae have not been investigated using cladistic methods: molecular data are available for less than a handful of species and morphological characters have not been coded and analyzed across the group. The Neotropical Leptophysoderes and Cryptophysoderes possess several characters thought to be plesiomorphic among Physoderinae and were suggested to be the putative sister-group of all remaining Physoderinae (Weirauch 2003), but this hypothesis remains to be tested. The aptery and resulting modifi ed morphology somewhat hamper the testing of hypotheses for the phylogenetic placement of the Afrotropical Porcelloderes based on morphology. Finally, it also remains to be tested whether the Madagascan Physoderinae are monophyletic, representing a single clade and potentially therefore a radiation in adaptation to different habitats or other biotic or abiotic factors. Alternatively, the Madagascan fauna might represent several unrelated lineages, as recently shown for the assassin bug subfamily Ectrichodiinae (Forthman & Weirauch 2016).
Even though some of the smaller genera are likely monophyletic, this is not necessarily the case for Physoderes: examination of specimens from various institutions, including types, has revealed undocumented morphological diversity that may challenge the monophyly of this genus. Physoderes currently contains 38 species mostly described by Miller (e.g., 1940Miller (e.g., , 1941, with additional species contributed by Westwood (1847), Stål (1863Stål ( , 1870, Horváth (1900), Breddin (1903), Kirkaldy (1905), Bergroth (1906), Distant (1903Distant ( , 1909, China (1935), Usinger (1946), Cao et al. (2011) and Davranoglou (2014). Physoderes has never been taxonomically revised, almost all descriptions lack proper diagnoses as well as high-quality habitus and genitalic illustrations, and no identifi cation key for the genus is in existence, resulting in a large number of misidentifi ed museum specimens. In addition, many species descriptions are based on singletons, with either males or females assigned as holotypes, which can be problematic due to the presence of sexual dimorphism in some species. A sizeable number of Physoderes specimens have accumulated in various museums (we have located 905 specimens in 13 natural history collections) and we believe that it is timely to produce a modern monograph of the Oriental/Australasian fauna of Physoderinae, while revising generic concepts across the group and recognizing monophyletic groups as genera.

Fig. 2.
Phylogeny of Physoderinae based on 57 morphological characters for 57 taxa analyzed using parsimony on TNT. Open white circles denote homoplastic characters, black circles denote characters that are not homoplastic. Numbers above the circles refer to character number (Table 1) and numbers below the circles refer to the character states (Table 3). Numbers in bold indicate symmetric resampling values.

Taxon sampling
A total of 57 taxa (3 outgroup taxa: Aradomorpha, Microlestria, Nalata; 54 ingroup taxa) were examined. All 14 genera of Physoderinae were sampled (Fig. 3), with all species represented except in Physoderes   Fig. 3. Dorsal habitus images of other specimens examined for morphological coding. Images by Jean-Michel Berenger, except Leptophysoderes sarapiqui Davranoglou, Hwang & Weirauch, 2015 and Aradomorpha crassipes Champion, 1899. Scale bars = 2 mm.  Miller, 1955a (5 of 6 sampled). Species of Physoderes were selected to represent what we perceive as morphogroups that may not be monophyletic and therefore were important to include in this analysis. Ten newly described species are also included to determine their genus-level placement. Whenever the association between males and females was in doubt, such as when the type specimen and its allotype are morphologically different, both were included in the analysis.

Morphological dataset
Type specimens for all species were examined at the British Museum of Natural History (BMNH) and the Muséum national d'Histoire naturelle (MNHN). Additional specimens were loaned from different institutions and examined at the University of California, Riverside (UCR) using a Nikon SMZ1500 stereo microscope and recording morphological measurements. Scoring of characters was based on type specimens, plus additional identifi ed material whenever available. A total of 57 morphological characters were coded using MESQUITE (Maddison & Maddison 2011;Tables 1-2), 50 as discrete characters that are either binary (24 characters) or unordered multistate (26 characters). A total of 7 continuous characters based on ratios of body measurements were also included ( Table 2).

Analysis parameters
A cladistic analysis was performed using a TNT (Goloboff et al. 2008) new technology search with initial level set at 50, 80 and 100, fi nding minimum length 10 times, and otherwise default settings. Symmetric resampling was performed using default settings for 500 replicates. Optimization of morphological characters on the resulting most parsimonious tree was performed in WinClada (Nixon 2002).

Specimens examined
A total of 905 specimens were examined during the course of this study. The specimens were loaned from various museums (see list of museum acronyms below). Holotype specimens described by Miller   information for each species was generated from the database and provided the basis for the "material examined" section as part of each species description.
Almost all locality labels lacked geographic coordinates and were therefore subsequently georeferenced using Google Earth. Localities with generic provincial or country information were georeferenced to the center of the least inclusive geographic region provided. Distribution maps were generated based on the georeferenced data and created using the AMNH Simple Mapper online tool linked to the PBI database. The maps are organized according to genus, except for Physoderes, which is split into two maps to avoid heavy overlap of specimen points.

Morphological methods
Habitus images were produced using either a Microptics-USA imaging system with a Canon EOS 1D camera or an Automontage GT-Vision imaging system. Images are either single exposures or combined from raw images using the Helicon Focus version 4.16 software or Archimed. To remove male genitalia, specimens were relaxed overnight in a hot water bath before soaking the apex of the abdomen in 10% potassium hydroxide (KOH) for 2 hours. Male genitalia were dissected after being heated in a 10% KOH solution for 8 minutes, neutralized in water and transferred into 99% glycerol on a ceramic spot plate. The endosoma remained contracted for all specimens examined as we were unable to completely infl ate it. The pygophores, parameres and phallus were dissected and mounted in a glycerin-gelatin mixture for imaging with an Automontage GT-Vision imaging system. Dissected genitalia were stored

Results of the phylogenetic analysis
The morphological coding for each taxon is shown in Table 1, measurements used as continuous characters in Table 2, and the list of the morphological character states used in Table 3. The phylogenetic analysis consistently produced a single most parsimonious tree (length = 365, CI = 0.22, RI = 0.61, Fig. 2; symmetric resampling values greater than 50 reported) that identifi ed the Neotropical Aradomorpha as the sister-group to Physoderinae, the Afrotropical Porcelloderes as sister to all remaining Physoderinae and a monophyletic New World clade containing Leptophysoderes and Cryptophysoderes. The Madagascan Physoderinae are separated into at least two clades and are therefore not monophyletic (Fig. 2). Physoderes mauriciensis (Villiers, 1964)    European Journal of Taxonomy 341: 1-118 (2017) 14 oddly dorso-ventrally fl attened Mimoelasmodema depressa Villiers, 1962, the other containing all other documented Madagascan Physoderinae (Fig. 2). The two largest Madagascan genera, Neophysoderes and Epiroderoides, are not monophyletic; Epiroderoides is rendered paraphyletic by the remaining Madagascan physoderine genera included in Clade B. Similarly, our analysis indicates that Physoderes is polyphyletic, such that species that fall into three distinct clades and one single-species lineage are currently classifi ed in this genus. We here propose a revised classifi cation that splits Physoderes into four genera, describing three of them as new: Nanophysoderes gen. nov. for N. dentiscutum (Bergroth, 1906) comb. nov. from Indonesia, which is recovered as the sister species to all Oriental, Australasian, and Madagascan Physoderinae; Macrophysoderes gen. nov. for a clade currently comprised of eight previously described or new species with distributions ranging from peninsular Malaysia to Papua New Guinea; and Breviphysoderes gen. nov., also with eight species that are widely distributed in Southeast Asia. We refrain from re-classifying the two non-monophyletic Madagascan genera, pending additional character investigation.
The overall morphology of Physoderinae is rather conserved and identifying phylogenetically informative characters proved challenging. Most of the characters that vary between species are found on the head and pronotum. However, our analysis shows that many of these features tend to be homoplastic, either having evolved multiple times across the phylogeny or having been lost on some members of a clade.

Diagnosis
Recognized among all other genera of Physoderinae by the head being elongate conical, without a pair of long, straight setae on the posterior margin of the postocular lobe; the scape surpassing the apex of the clypeus; the eye being distinctly projecting in dorsal view; the pronotum with setigerous tubercles bearing short, curved setae; the anterior pronotal lobe always narrower than the posterior lobe in both sexes; the parascutellar lobe being semicircular; the posterior margin of the connexivum often being slightly elevated, with short, curved setae; the anterior margin of the male mediosternite 8 sharply emarginated; the cup-like sclerite of the pygophore apically rounded, with paired sclerotized latero-ventral slight protuberances; the arms of the basal plate parallel to each other; the sclerotized ring of the ductifer elongate; the dorsal phallothecal sclerite subacute, with lateral narrow; and the bandlike prolongations of the dorsal phallothecal sclerite extended diagonally towards the basal plate. Most similar to Physoderes, but distinguished by the scape surpassing the apex of the clypeus and the anterior pronotal lobe being narrower than the posterior lobe in males.

Etymology
Named after Physoderes, the type genus of the subfamily Physoderinae, in combination with "brevi" (Latin for "short") to indicate the shorter body length. The gender is feminine.
HEAD. Elongate conical; maxillary plate truncate apically; scape surpassing apex of clypeus except in Breviphysoderes decora comb. nov.; eye distinctly projecting in dorsal view except in B. decora comb. nov., not attaining head ventral margin in lateral view; height of anteocular lobe shorter than postocular lobe, ocelli present.
THORAX. Antero-lateral paired projections acute, oriented anteriorly or diverging; surface of anterior lobe with low ridges; median pronotal depression contiguous with transverse sulcus in males except in Breviphysoderes vestita comb. nov.; paramedian carina strongly defi ned except in B. decora comb. nov.; posterior lobe medially rugose; anterior pronotal lobe of equal length to posterior lobe, narrower than posterior lobe in both sexes, anterior lobe lower than posterior lobe in lateral view except in B. vestita comb. nov.; parascutellar lobe semicircular; scutellum rounded triangular, scutellar process long, with rounded apex, except in B. decora comb. nov.; mesosternite usually with median irregular tuberculated protrusion between fore and mid coxae.
ABDOMEN. Elongate ovoid, with rounded or straight terminal margin; connexival margin slightly undulating except in Breviphysoderes planicollis comb. nov., posterior margin most often slightly elevated.
GENITALIA. Anterior margin of mediosternite 8 sharply emarginate, with or without medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, with apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded, with sclerotized paired latero-ventral slight protuberance; arms of basal plate parallel to each other; ductifer with sclerotized, elongate ring; endosomal struts conical, with subacute apex, basally divided into two arms; apex of dorsal phallothecal sclerite subacute or rounded, with lateral narrow band-like prolongations oriented diagonally towards basal plate.

Female
Similar in shape and coloration to males but slightly larger or having a wider abdomen or narrower anterior pronotal lobe.

Ecology
Specimens have been collected from dead plant material, tree bark and in the undergrowth. Different species have been found at different elevations with B. vestita comb. nov. collected at the highest altitude of 1955 m. hemelytron attaining the apex of the abdomen, and the anterior half of the connexivum brown and the posterior half straw-colored. It most closely resembles B. mjoebergi (Miller 1940) comb. nov., but can be differentiated by the scape not extending beyond the clypeal apex and the shorter scutellar process for B. mjoebergi comb. nov.   COLORATION . Light brown to dark brown. Head light brown to dark brown. Scape of antenna straw-colored, with apex brown, pedicel straw-colored and medially brown, basifl agellomere dark brown, distifl agellomere dark brown with straw-colored apex. First segment of labium brown or dark brown, second segment light brown and basally and apically brown or dark brown entirely, third segment brown or dark brown. Anterior lobe of pronotum dark brown with straw-colored ridges, posterior lobe light brown or brown, scutellum dark brown with straw-colored apex, pleuron dark brown entirely or with brown suffusion, sternum dark brown. Corium of hemelytron brown, membrane brown with apical veins of external cell (Cu + M) straw-colored. Femora of legs straw-colored or light brown with medial and apical dark brown annulations, tibiae straw-colored or brown with basal, medial and apical dark brown annulations, tarsi and claws light-brown or brown. Abdomen not examined dorsally, ventrally light brown medially with lateral suffusion to dark brown or entirely dark brown, anterior half of connexivum dark brown and posterior half straw-colored, or dark brown entirely, exposed part of pygophore variable, light brown to dark brown.

Material
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with sparse, setigerous tubercles, without 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Fore femur of legs with two rows of spines and setigerous tubercles, fore tibia with regular rows of tuberculated, stout, sharp setae. On abdomen, posterior margin of connexivum with short, curved setae.
HEAD. Scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view.
HEMELYTRON. Not attaining tip of abdomen.
LEGS. Same as genus description.

Ecology
This species is found in the tropical forest of Borneo and has been collected from the undergrowth, sapwood just under bark, and fl ood refuse and cut reeds at the junction of rivers.

Distribution
This species is known only from the type locality of Mt Dulit in Sarawak, north-central Borneo. All three specimens were collected near each other, at the foot of Mt Dulit near the River Tinjar.

Remarks
This species is described based on three specimens originally designated as paratypes of B. hobbyi (Miller, 1940) comb. nov. Even though they share the same collecting event and locality as that of B. hobbyi, closer examination shows distinct, consistent, morphological differences and the specimens are therefore described as a separate species. The specimens were collected by B.M. Hobby and A.W. Moore during the Oxford University Expedition in 1932. The darkened scutellum and overlapping distribution shared with B. hobbyi comb. nov., B. decora comb. nov., B. planicollis comb. nov. and B. tenebrosa gen. et sp. nov. suggest that they are likely to be closely related. (Miller, 1940) comb. nov.

Diagnosis
This species is recognized among species of Breviphysoderes by the head and anterior pronotal lobe being distinctly covered with dense, short, curved, setigerous tubercles, the deeply defi ned pronotal paramedian carina, the hemelytron surpassing the tip of the abdomen, the medial vein of the hemelytron shaped like an inverted S, and the elongated external cell of the hemelytron (Cu + M). This species is most similar to B. tenebrosa gen. et sp. nov. and B. planicollis comb. nov. It can be differentiated from the former by the overall coloration, the diverging antero-lateral pronotal projections, and the shape of the external cell. It differs from B. planicollis comb. nov. by the overall coloration and the shape of the external cell.

Redescription
Male BODY LENGTH. Medium size, total length 8.7 mm. . Light brown with dark brown patterns. Head brown. Scape of antenna strawcolored with light brown apex, basal half of pedicel straw-colored and apical half brown, basifl agellomere brown and distifl agellomere brown with light brown apex. Labium brown. Anterior lobe of pronotum dark brown, posterior lobe brown, scutellum dark brown with brown apex, pleuron dark brown with brown suffusion. Corium and membrane of hemelytron brown. Femora of legs straw-colored with medial dark brown annulation and apical dark brown suffusion, tibiae straw-colored with basal, medial and apical dark brown annulations, tarsi and claws straw-colored. Abdomen not examined dorsally, European Journal of Taxonomy 341: 1-118 (2017) 24 ventrally light brown medially with lateral suffusion to dark brown, connexivum dark brown with apex straw-colored, exposed part of pygophore brown.
VESTITURE. Sparsely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with sparse, setigerous tubercles, without 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum with short, curved setae.
HEAD. Scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view.
LEGS. Same as genus description.

Ecology
This species is found in old secondary forest of Borneo.

Distribution
This species is only known from the type locality of Mt Dulit in Sarawak, north-central Borneo. The type locality is shared with B. fulvopicta gen. et sp. nov., B. planicollis comb. nov., B. decora comb. nov. and B. tenebrosa gen. et sp. nov.

Remarks
This species is redescribed based only on the holotype specimen. Originally, fi ve paratype specimens were associated with the holotype from the same area. Closer examination of these specimens showed that they are not conspecifi c. We here treat these specimens as the two newly described species B. fulvopicta gen. et sp. nov. and B. tenebrosa gen. et sp. nov., and also as B. decora comb. nov. Breviphysoderes hobbyi comb. nov. is removed from Physoderes here and transferred to the new genus because it possesses its synapomorphies (semicircular parascutellar lobes, males do not possess an infl ated anterior pronotal lobe and have prominent setigerous tubercles on the dorsal ridges of the anterior pronotal lobe). The specimen was collected by B.M. Hobby and A.W. Moore during the Oxford University Expedition in 1932. (Miller, 1940), comb. nov.

Diagnosis
Recognized among species of Breviphysoderes gen. nov. by the smaller size, the head and anterior pronotal lobe being distinctly covered with dense, short, curved, setigerous tubercles, the deeply defi ned pronotal paramedian carina, the slightly elevated and setose posterior margin of the connexivum and the ductifer having a sclerotized elongate ring. This species is most similar to B. vestita comb. nov. but can be differentiated by its smaller size, the lack of an infl ated anterior pronotal lobe in males, the hemelytron not surpassing the tip of the abdomen, and the slightly undulating (vs angularly hooked), sclerotized, fl ap-like prolongations of the phallosoma.
COLORATION . Straw-colored with dark brown patterns. Head straw-colored to light brown with dark brown suffusion. Scape and pedicel of antenna straw-colored to light brown, with slight apical darkening, basifl agellomere distinctly darker than scape and pedicel, light brown to dark brown, distifl agellomere straw-colored to light brown with base dark brown. First segment of labium strawcolored to brown, second segment basally and apically dark brown, medially straw-colored or light brown, third segment dark brown. Anterior lobe of pronotum dark brown with straw-colored ridges, posterior lobe straw-colored or light brown, scutellum basally dark brown, progressively lightening towards a straw-colored apex, pleuron straw-colored with dark brown suffusion, sternum straw-colored with dark brown suffusion. Corium of hemelytron straw-colored or light brown, membrane dark brown. Femora of legs straw-colored with medial and apical dark brown annulations, tibiae straw-colored with basal, medial and apical dark brown annulations, tarsi and claws straw-colored. Abdomen dorsally yellowish-orange, ventrally brown with straw-colored lateral margins, sub-lateral and medial spots, and dark brown sub-lateral suffusion, anterior half of connexivum dark brown and posterior half strawcolored, with the color proportions slightly variable, exposed part of pygophore straw-colored.
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with sparse, setigerous tubercles, without 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum with short, curved setae. THORAX. Antero-lateral paired projections acute, diverging; median pronotal depression contiguous with transverse sulcus; paramedian carina strongly defi ned; scutellar process long, with rounded apex; mesosternite obscured.
LEGS. Same as genus description.

Female
Similar in shape and coloration as males, but some individuals slightly larger.

Ecology
Unknown.

Distribution
Vietnam, Borneo and Java. This species has a relatively wide distribution, with specimens collected from northern Vietnam and Java, which are near the northern-and southern-most limits of the distribution of Oriental physoderine species.

Remarks
Originally described as Physoderes mjoebergi by Miller (1940), this species is transferred to Breviphysoderes gen. nov. based on the phylogenetic analysis conducted above. It is most closely related to B. vestita (Horváth, 1900) comb. nov. and both species are found on the island of Java. The two female specimens from Vietnam are slightly larger than those from Borneo and Java, but do not have any differential morphological characters and are thus identifi ed as conspecifi cs. Physoderes dyak (Miller, 1955b) is synonymized here with B. mjoebergi comb. nov. based on the examination of one paratype at BMNH together with the holotype of B. mjoebergi comb. nov. No diagnostic characters were found to separate the two species based on the examination of these two specimens. Conspecifi city was further confi rmed by examination of habitus images of the P. dyak holotype at the ZMAN. (Miller, 1940)

Redescription
Male BODY LENGTH. Medium size, total length 9.8 mm.
COLORATION . Light brown with dark brown patterns. Head brown with dark brown suffusion. Scape of antenna straw-colored with brown apex, pedicel brown with straw brown apex, basi-and distifl agellomeres missing. First segment of labium brown, second segment basally and apically dark brown, medially light brown, third segment dark brown. Anterior lobe of pronotum dark brown with straw-colored patterns, posterior lobe straw brown, scutellum dark brown with brown apex, pleuron light brown with dark brown suffusion, sternum brown with dark brown suffusion. Corium of hemelytron straw-colored to brown, membrane straw brown to brown, slightly translucent. Femora of legs strawcolored with medial and apical light brown annulations, tibiae straw-colored with basal, medial and apical light brown annulations, tarsi and claws straw-colored. Abdomen dorsally yellowish-orange, ventrally brown with dark brown lateral suffusion.
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with sparse, setigerous tubercles, without 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, setigerous tubercles along lateral margins and sparsely distributed on 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. Posterior margin of connexivum with short, curved setae.
HEAD. Scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, less than 1/5 length of head, not attaining ventral margin of head in lateral view.
LEGS. Same as genus description.
GENITALIA. Previously dissected and genitalia missing.

Female
Unknown.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is only known from the type locality of Mt Dulit in Sarawak, north-central Borneo. The type locality is shared with B. fulvopicta gen. et sp. nov., B. hobbyi comb. nov., B. decora comb. nov. and B. tenebrosa gen. et sp. nov.

Remarks
This species is redescribed based on the holotype specimen. Miller (1940) recognized this specimen as a species different from B. hobbyi (Miller, 1940) comb. nov. due to the different anterior pronotal lobe coloration. Breviphysoderes planicollis comb. nov. is removed from Physoderes and transferred to this genus because it shares its synapomorphies (parascutellar lobes are semicircular, the male specimen does not possess an infl ated anterior pronotal lobe and has prominent setigerous tubercles on the dorsal ridges of the anterior pronotal lobe). The specimen was collected by Dr E. Mjöberg during the Kalabit Expedition. The darkened scutellum and overlapping distribution, shared with B. hobbyi comb. nov., B. decora comb. nov. and B. tenebrosa gen. et sp. nov., suggest that they are likely to be closely related. (Miller, 1940), comb. nov. Figs 4, 6;Appendix Physoderes shelfordi Miller, 1940: 551, fi g. 91.

Diagnosis
This species is recognized among other species of Breviphysoderes gen. nov. by the scape surpassing the apex of the clypeus, the antero-lateral pronotal projection oriented anteriorly, the deeply defi ned pronotal paramedian carina, the distinct straw-colored mark on the apical veins of the external cell of the hemelytron (Cu + M) and hemelytron surpassing the tip of the abdomen. It is most similar to B. hobbyi comb. nov. and B. planicollis comb. nov. It can be differentiated from B. tenebrosa gen. et sp. nov. by the overall coloration and a narrower postocular lobe. It differs from B. planicollis comb. nov. by the orientation of the antero-lateral pronotal projection and the straw-colored mark on the apical veins of the external cell.

Redescription
Male BODY LENGTH. Medium, total length 9 mm. COLORATION (Fig. 4). Light brown with dark brown patterns. Head light brown to dark brown. Scape of antenna straw-colored with light brown apex, basal half of pedicel straw-colored and apical half brown, basi-and distifl agellomeres missing. Labium light brown. Pronotum dark brown with brown suffusion, scutellum dark brown with light brown apex. Corium of hemelytron brown, membrane brown with apical veins of external cell (Cu + M) straw-colored. Femora of legs light brown with medial and apical dark brown annulations, tibiae brown with basal, medial and apical dark brown annulations, tarsi and claws light-brown or brown. Abdomen not examined dorsally, ventrally light brown medially with lateral brown suffusion, anterior half of connexivum dark brown and posterior half straw-colored.
VESTITURE. Sparsely setose. Head with some fl at, curved setae, without 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum with short, curved setae.
HEAD. Scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view.
LEGS. Same as genus description.

Female
Unknown.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is only known from the type locality in north-western Borneo, Kuching.

Remarks
This species is redescribed based on the holotype specimen. Breviphysoderes shelfordi comb. nov. is removed from Physoderes and transferred to Breviphysoderes gen. nov. because it possesses the synapomorphies of this genus (parascutellar lobes are semicircular, males do not possess an infl ated anterior pronotal lobe and have prominent setigerous tubercles on the dorsal ridges of the anterior pronotal lobe). This species is quite similar to other members of the genus found in Sarawak, Borneo, but is recognized as a separate species based on the diagnostic characters listed above. More specimens from this area will be needed to further test the status of these closely related species.

Diagnosis
This species is recognized among other species of Breviphysoderes gen. nov. by the entirely brownishblack coloration, the antero-lateral pronotal projection oriented anteriorly, the pronotal paramedian carina being deeply defi ned and the hemelytron surpassing the tip of the abdomen. This species is most similar to B. hobbyi comb. nov. and B. planicollis comb. nov. It can be differentiated from both species by the overall coloration, the orientation of the antero-lateral pronotal projection and the shape of the external cell.

Etymology
The name tenebrosa is selected after the Latin adjective for "tenebrosus", meaning dark, to describe the distinctive brownish-black coloration of the specimen.

Description
Male BODY LENGTH. Small size, total length 7.07 mm. . Entirely brownish-black. Antenna missing. Forelegs missing, mid-and hindlegs brownish black.
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with sparse, setigerous tubercles, without 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum with short, curved setae.
HEAD. Eye distinctly projecting in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view.

Female
Unknown.

Ecology
The type specimen was found on the bark of a felled tree along the junction of two rivers.

Distribution
This species is only known from the type locality of Mt Dulit in Sarawak, north-central Borneo. The type locality is shared with B. fulvopicta gen. et sp. nov., B. planicollis comb. nov., B. decora comb. nov. and B. hobbyi comb. nov.

Remarks
This species is described based on a single specimen originally designated as a paratype of B. hobbyi (Miller, 1940) comb. nov. Even though it shares the collecting event and locality with specimens of that species, closer examination showed distinct morphological differences and we describe this specimen as a separate species. It was collected by B.M. Hobby and A.W. Moore during the Oxford University Expedition in 1932. (Horváth, 1900) Physoderes vestita -Maldonado 1990: 381.

Diagnosis
This species is recognized among species of Breviphysoderes gen. nov. by having the head and anterior pronotal lobe distinctly covered with dense, short, curved, setigerous tubercles, the scape surpassing the apex of the clypeus, males with the anterior pronotal lobe almost as wide as the posterior pronotal lobe, the apical veins of the external cell of the hemelytron (Cu + M) with distinct straw-colored marks, and the hemelytron surpassing the tip of the abdomen. This species most closely resembles B. fulvopicta gen. et sp. nov. but is differentiated by the wider anterior pronotal lobe in males and the length of the hemelytron (i.e., surpassing the tip of the abdomen).

Redescription
Male BODY LENGTH. Medium-large size, total length 10.02 mm, SD ± 0.24 (Appendix). 7). Straw-colored to brown with dark brown patterns. Head brown. Scape of antenna straw-colored with light brown apex, pedicel straw-colored to light brown, basifl agellomere straw-colored with light brown apex, distifl agellomere straw-colored with light brown base. First segment of labium with basal half brown, apical half straw-colored, second segment straw-colored with base and apex brown, third segment brown. Pronotum dark brown with straw-colored setae, scutellum dark brown, pleuron straw-colored to dark brown, sternum brown to dark brown. Corium of hemelytron straw-colored, membrane brown with apical veins of external cell (Cu + M) straw-colored, sometimes including medial vein apex. Femora straw-colored with medial and apical brown annulations, tibiae straw-colored with basal, medial and apical light brown annulations, tarsi and claws straw-colored. Abdomen dorsally yellowish-orange, ventrally light brown, medially with dark brown sub-lateral and posterior suffusion, anterior half of connexivum dark brown and posterior half straw-colored, exposed part of pygophore straw-colored.
VESTITURE. Densely setose. Head with widespread setigerous tubercles bearing long, semi-erect, spatulate, curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, with a 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, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum with short, curved setae.
HEAD. Scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe with raised ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina strongly defi ned; scutellar process long, with rounded apex; mesosternite with median irregular tuberculated protrusion between fore and mid coxae.
LEGS. Same as genus description.

Female
Similar to males except in having a narrower anterior pronotal lobe.

Ecology
Nothing is known about the biology of this species except that specimens have been collected from mid to high altitudes ranging from 447 m to 1955 m with most specimens collected from elevations higher than 1200 m.

Distribution
This species is exclusively found on the island of Java (the locality for one of the examined specimens is unknown).

Remarks
Originally described as Epirodera vestita (Horváth, 1900), this species is transferred to Breviphysoderes gen. nov. based on the phylogenetic analysis conducted above. It is most similar to B. mjoebergi (Miller, 1940) comb. nov., and both species co-occur on the island of Java. Physoderes serraticollis Breddin, 1903, P. javanica Miller, 1940and P. rugosa Miller, 1954 are here synonymized with B. vestita comb. nov. based on the examination of all type specimens. No diagnostic morphological characters can be found to separate these species, except that the holotype of P. javanica, which is a female, has a narrower anterior pronotal lobe. Nevertheless, this specimen possesses the raised anterior pronotal ridges with setigerous tubercles, the completely dark brown anterior pronotal lobe, the straw-colored apical veins of the external cell of the hemelytron and the rounded abdominal margin that are diagnostic for B. vestita comb. nov. The holotypes of both P. rugosa and P. serraticollis are males and share the broad anterior pronotal widths of B. vestita, which has raised ridges and dense, tuberculated, curved setae, as well as the remaining diagnostic characters. Both were also collected from Java. , the eyes being very large and distinctly projecting in dorsal view and sometimes attaining the ventral margin of the head in lateral view, the anterior pronotal lobe narrower or slightly narrower than the posterior lobe, the antero-lateral pronotal projection often being truncate, but sometimes acute, and the margin of the anterior opening of the pygophore being rounded in lateral view. Macrophysoderes gen. nov. is most similar to Physoderes, but is distinguished by the very large and distinctly projecting eyes in most species, the males with the anterior pronotal lobe narrower than the posterior lobe and by the glabrous pronotum.

Etymology
The name combines Physoderes after the type genus of Physoderinae and the Greek adjective "macro" to indicate the large size of the eyes. The gender is feminine.
HEAD. Elongate or short conical; maxillary plate truncate or rounded apically; scape length variable; eye distinctly projecting in dorsal view except in M. elongata gen. et sp. nov., sometimes attaining ventral margin of head in lateral view; height of anteocular lobe shorter or level with postocular lobe, ocelli present.
THORAX. Antero-lateral paired pronotal projection truncate or acute, oriented anteriorly or diverging; surface of anterior lobe smooth, anterior pronotal ridges almost obsolete or low; median pronotal depression contiguous with transverse sulcus in both sexes except in M. cirripilosa gen. et sp. nov. and M. elongata gen. et sp. nov.; paramedian carina variably defi ned; posterior lobe medially rugose; anterior pronotal lobe equal to or shorter than posterior lobe, narrower or slightly narrower than posterior lobe in males, anterior lobe lower than or level with posterior lobe in lateral view; parascutellar lobe shape variable, either triangular, bell-shaped or rounded and skewed towards median; scutellum rounded triangular, length of scutellar process variable from very short to long, shape of apex variable; mesosternite with median, irregular, tuberculated protrusion between fore and mid coxae except in M. elongata gen. et sp. nov.
ABDOMEN. Elongate ovoid, with straight or rounded posterior margin; connexival margin smooth or slightly undulating, posterior margin not elevated.
GENITALIA. Anterior margin of mediosternite 8 undulating or sharply emarginated, with or without medial apodeme; transverse bridge of pygophore of variable width, margin of anterior opening rounded or angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically variable; arms of basal plate converging or curved to form rounded foramen; ductifer with membranous or sclerotized ring of variable shapes; endosomal struts of variable shapes; dorsal phallothecal sclerite subacute, with lateral, broad, plate-like prolongations of variable sizes.

Female
Most often similar in size, shape and color to male except with wider abdomen.

Ecology
Not much is known about the biology. A few species are known to occur at higher elevations (1000-2000 m). One specimen of M. cirripilosa gen. et sp. nov. was collected from a log and another from a light trap.

Distribution
This genus currently comprises 8 species and is widely distributed across Southeast Asia and Australasia, and can be found in the Bengal region, peninsular Malaysia, Borneo, New Guinea and the Solomon Islands. The highest species diversity for this genus is found on the island of New Guinea.

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the dark brown coloration, the scape reaching the apex of the clypeus, the very large eyes relative to the head, the absence of paired long setae at the posterior margin of the postocular lobe, the two rows of small setigerous tubercles on the ventral surface of the postocular lobe, the pronotum with deep paramedian carinae, the triangular pronotal paramedian lobes, the spines on the fore trochanter and the straight margin of the abdominal apex. This species is quite distinct from other members of the genus.

Redescription
Male BODY LENGTH. Medium size, total length 9 mm. COLORATION (Fig. 10). Brown. Head brown. Scape of antenna brown, other segments missing. First segment of labium brown, second and third straw brown. Anterior lobe of pronotum dark brown, posterior lobe brown, scutellum dark brown, pleuron brown with light brown margin along acetabulum, sternum brown. Corium of hemelytron brown with straw brown apex, membrane dark brown. Femora straw brown with sub-basal and sub-apical brown annulations, tibiae brown, basally straw brown, tarsi and claws straw brown. Abdomen dorsally straw brown, ventrally straw brown with lateral brown patterns, connexivum yellow brown with anterior indistinct brown spot, exposed part of pygophore straw-colored.
VESTITURE. Sparsely setose. Head with some fl at, curved setae, ventral surface of postocular lobe with two rows of small, setigerous tubercles. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, posterior lobe with only short, sparse setae. Corium of hemelytron glabrous. Fore trochanter of legs with small process on internal surface, fore femur with two rows of spines and setigerous tubercles, fore tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen without setae. HEAD. Short conical; maxillary plate rounded apically; scape reaching apex of clypeus; eye about 1/5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe shorter than postocular lobe.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe with low ridges; paramedian carina strongly defi ned; posterior lobe medially weakly rugose; anterior pronotal lobe shorter than posterior lobe, narrower than posterior lobe, anterior lobe lower than posterior lobe in lateral view; parascutellar lobe triangular; scutellar process long, with acute apex.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore extremely narrow, only consisting of margin of anterior opening of pygophore, margin of anterior opening rounded; cup-like sclerite apically rounded, with sclerotized paired latero-ventral protuberance; arms of basal plate converging; ductifer with sclerotized elongate ring; endosomal struts apically truncate, divided into two arms; shape of dorsal phallothecal sclerite subacute with lateral broad, plate-like prolongations, small acute apex and broad lateral plates with rounded margins extended laterally and curved towards apex.

Female
Unknown.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is only known from two localities: Pusa, Bengal (type locality) and Khasia Hills, Assam. It appears to be restricted to areas adjoining the Indian subcontinent and Myanmar.

Remarks
This species is placed in the Macrophysoderes gen. nov. clade based on the phylogenetic analysis above. It is most closely related to M. monticola comb. nov., as both possess a scape that reaches the apex of the clypeus and lack the paired, long setae at the posterior margin of the postocular lobe. M. bengalensis comb. nov. possesses several unique characters, especially on the male genitalia.

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the short conical head that is covered with semi-erect, curved setae, with a distinct concentration of setae on the postocular lobe between the ocelli, the length of the scape reaching the apex of the clypeus, the very large eye that attains the ventral margin of the head in lateral view, the median pronotal depression not contiguous with the transverse sulcus and the short scutellar process. It most closely resembles M. fi nisterre gen. et sp. nov. and M. elongata gen. et sp. nov. It is differentiated from the former by the overall coloration, the shape of the antero-lateral pronotal projections, the wider anterior pronotal lobe and the shape of the abdominal apical margin. It is differentiated from M. elongata gen. et sp. nov. by the size, the shape of the scutellar process, the relative length of head to body and the less swollen fore femur.

Etymology
The name combines the Latin noun "cirrus", meaning curl or hair-tuft, with the Latin adjective "pilosus", meaning hairy, to describe the abundant, curved setae on the head, especially between the ocelli.
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with two rows of small, 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, tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen with a few clubbed setae on each segment. HEAD. Short conical; maxillary plate rounded apically; scape reaching apex of clypeus; eye distinctly projecting in dorsal view, about 1/ 5 length of head, attaining ventral margin of head in lateral view; height of anteocular lobe shorter than postocular lobe.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe with low ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina strongly defi ned; posterior lobe medially weakly rugose; anterior pronotal lobe shorter than posterior lobe, slightly narrower than posterior lobe, anterior lobe lower than posterior lobe in lateral view; parascutellar lobe bell-shaped and skewed towards median; scutellar process short with rounded apex.
HEMELYTRON. Not attaining tip of abdomen.
LEGS. Same as genus description.

Female
Similar in size and shape to males except with rounder or wider abdomen. Fig. 11. Ventral habitus images for species of Macrophysoderes gen. nov. and Nanophysoderes gen. nov. Scale bars = 2 mm.

Ecology
One specimen was recorded as having been found in logs and another as caught in a light trap.

Distribution
Found throughout the island of New Guinea, on the nearby island New Britain, and on Bougainville Island (Solomon Archipelago).

Remarks
This species is described based on the examination of 147 specimens. It is the most commonly collected physoderine from the island of New Guinea. There are slight differences among specimens collected from different regions, most apparently variation of size and color. These differences include overlapping variation and are therefore not indicative of multiple species. Genitalic dissections of specimens from different regions showed no differences. Based on the phylogenetic analysis above, the species is placed in the newly created

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the large size, the short conical head with semi-erect, curved setae on the postocular lobe between the ocelli, the length of the scape reaching the apex of the clypeus, the very large eye that does not attain the ventral margin of the head in lateral view, the acutely diverging antero-lateral pronotal projections, the wide anterior pronotal lobe that is slightly narrower than the posterior pronotal width, the short and spatulate scutellar process, the extremely incrassate fore femur, the hemelytron surpassing the abdominal tip and the abdominal terminal margin being straight. This species is closest to M. grandis gen. et sp. nov., but is differentiated by the body size, head shape, scape length, eye shape and the shape of the anterior pronotal lobe.

Etymology
The name elongata is an adjective based on the Latin participle "elongatus" meaning elongate to describe the elongated abdomen unique to this species. COLORATION (Fig. 10). Brown. Head brown. Scape of antenna straw-colored to light brown, sometimes with brown apex, pedicel basally brown and apically straw-colored, basifl agellomere brown, distifl agellomere basally brown, apically straw-colored. First segment of labium basally brown, apically light brown, second segment basally and apically brown, medially straw-colored or light brown, third segment brown. Anterior lobe of pronotum dark brown with light brown markings, posterior lobe brown, scutellum basally dark brown, apically including process light brown, pleuron with mixture of strawcolor to dark brown, sternum dark brown. Corium of hemelytron brown to dark brown, membrane dark brown. Femora of legs light-brown with sub-basal and apical dark brown annulations, tibiae light brown with basal, medial and apical dark brown annulations, tarsi and claws brown. Abdomen dorsally yellowish orange, ventrally brown medially with sub-lateral dark and light brown patterns, anterior half of connexivum dark brown, posterior half light brown, exposed part of pygophore brown.

Holotype
VESTITURE. Sparsely setose. Head with widespread curved setae, ventral surface of postocular lobe with two rows of small, setigerous tubercles, with pair of long, straight setae on postocular lobe, posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, posterior lobe with short, curved, setigerous tubercles along lateral margins and sparsely distributed on dorsal surface. Corium of hemelytron with short, curved setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen with no prominent paired setae, or connexival margin with a few clubbed setae on each segment. HEAD. Short conical; maxillary plate rounded apically; scape reaching apex of clypeus, or just surpassing apex of clypeus; eye hemispherical in dorsal view, about 1/5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe shorter than postocular lobe.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe smooth, ridges almost obsolete; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defi ned; posterior lobe medially weakly rugose; anterior pronotal lobe of equal length to posterior lobe, slightly narrower than posterior lobe, anterior lobe level with posterior lobe in lateral view; parascutellar lobe bell-shaped and skewed towards median; scutellar process short, apex spatulate; mesosternite with setigerous tubercles, but no protrusion.

Female
Similar in size and coloration to males, with median depression of anterior pronotal lobe contiguous with transverse sulcus.

Ecology
This species was collected from high altitudes ranging between 1200 and 2100 m.

Distribution
This species is known to occur in the highlands of eastern Papua New Guinea.

Remarks
This species shares characters with M. cirripilosa gen. et sp. nov., including the setose, short, and conical head, the very large eyes, the shape of the parascutellar lobe and the anterior pronotal lobe being almost as wide as the posterior lobe.

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the overall brownish-black color, the short conical head with semi-erect, curved setae on the postocular lobe between the ocelli, the length of the scape that just surpasses the apex of the clypeus, the very large eye that attains the ventral margin of the head in lateral view, the very short antero-lateral pronotal projections that are acute and diverging, the narrow anterior pronotal lobe, the very short scutellar process, the narrow connexivum, and the straight abdominal terminal margin. It most closely resembles M. cirripilosa gen. et sp. nov. and M. elongata gen. et sp. nov. It is differentiated from the former by the overall coloration, the shape of the antero-lateral pronotal projections, the narrower anterior pronotal lobe and the shape of the abdominal apical margin. It differs from the latter by the size, the shape of the scutellar process, the relative length of the head versus the body and the only slightly swollen fore femur.

Etymology
The name fi nisterre is a noun in apposition named after the holotype locality, the Finisterre mountain range in Papua New Guinea.

Redescription
Male BODY LENGTH. Medium size, total length 8.85 mm, SD ± 0.38 (Appendix). COLORATION (Fig. 10). Brownish-black. Head brownish-black. Scape of antenna straw-colored to light brown with brown apex, pedicel light brown suffused with brown, basifl agellomere brown, distifl agellomere basally brown, apically straw-colored. First segment of labium light brown, second segment basally and apically brown, medially straw-colored or light brown, third segment brown. Pronotum brownish-black, with faint brown ridges, scutellum basally brownish-black, apically including process brown, pleuron mixture of straw-color to dark brown, sternum brownish-black. Corium of hemelytron brownish black, membrane brownish black. Femora light-brown with sub-basal and apical dark brown annulations, tibiae light brown with basal, medial and apical dark brown annulations, tarsi and claws brown. Abdomen dorsally dull yellow, ventrally straw-colored or light brown with dark brown suffusion or patterns, anterior half of connexivum dark brown, posterior half light brown, sometimes indistinct, exposed part of pygophore brown.
VESTITURE. Densely setose. Head with widespread curved, setigerous tubercles, ventral surface of postocular lobe with two rows of small, 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. HEAD. Short conical; maxillary plate rounded apically; scape reaching apex of clypeus or surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, attaining ventral margin of head in lateral view; height of anteocular lobe shorter than that of postocular lobe.
THORAX. Antero-lateral paired projections slightly acute, diverging or obsolete; surface of anterior lobe with low ridges; median pronotal depression contiguous with transverse sulcus; paramedian carina strongly defi ned; posterior lobe medially rugose; anterior pronotal lobe of equal length to posterior lobe, narrower than posterior lobe, anterior lobe level with posterior lobe in lateral view; parascutellar lobe bell-shaped and skewed towards median; scutellar process very short, with subacute apex.
LEGS. Same as genus description.

Female
Similar in size and shape to males except having a wider abdomen and connexivum.

Ecology
This species was collected from an altitude between 1000 and 1200 m.

Distribution
This species is known only from the Finisterre and Herzog mountain ranges in eastern Papua New Guinea.

Remarks
Based on the phylogenetic analysis above, this species forms a monophyletic clade together with two other species from Papua New Guinea within Macrophysoderes gen. nov.

Diagnosis
This species is recognized among other species of Macrophysoderes gen. nov. by its very large size, the elongate conical head, the scape surpassing the apex of the clypeus, the bulbous eye that reaches the ventral margin of the head in lateral view, the truncate antero-lateral pronotal projection, the anterior pronotal lobe being distinctly narrower than the posterior lobe, and the short and spatulate scutellar process. This species is distinctive based on these characters, especially its large size.

Etymology
The name grandis is an adjective after the Latin "grandis", meaning large, to describe the exceptionally large size of this species. Paratype PAPUA NEW GUINEA: 1 ♀, same collection data as holotype but 28 Aug. 1956 (UCR_ENT 00073625) (BPBM).

Description
Male BODY LENGTH. Very large size, total length 11.58 mm. . Brown with straw-colored markings. Head brown. Scape and pedicel of antenna straw-colored with brown suffusion, basifl agellomere brown, distifl agellomere straw-colored with brown base. First segment of labium light brown, second segment basally and apically brown, medially straw-colored, third segment brown. Pronotum brown, scutellum basally brown to dark brown, apically straw-colored or light brown, pleuron brown, sternum brown and straw-colored. Corium and membrane of hemelytron brown. Femora of legs straw-colored with sub-basal and apical brown annulations, tibiae straw-colored with basal, medial and apical brown annulations, tarsi and claws strawcolored. Abdomen dorsally dull yellow, ventrally light brown medially with brown and straw-colored sub-lateral patterns, anterior half of connexivum dark brown, posterior half straw-colored.
VESTITURE. Glabrous. Head with some fl at, curved setae, ventral surface of postocular lobe with a few curved setae, with pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and glabrous dorsal surface, posterior lobe glabrous. Corium of hemelytron glabrous. Legs with two rows of spines and setigerous tubercles, tibiae with a few prominent tuberculated, stout, sharp setae. Connexival margin of abdomen glabrous. HEAD. Elongate conical; maxillary plate rounded apically; scape surpassing apex of clypeus; eye distinctly projecting in dorsal view, about 1/5 length of head, reaching ventral margin of head in lateral view; height of anteocular lobe shorter than postocular lobe.

Female
Similar in size, shape and color to male, except with lighter colored scutellum.

Ecology
This species is only known from two specimens caught in a light trap set at an altitude of 1010 m.

Distribution
This species is known only from the type locality of Bulolo, northeast Papua New Guinea.

Remarks
This species shares characters with M. cirripilosa gen. et sp. nov., including the setose conical head, the very large eyes and the shape of the parascutellar lobe. (Miller, 1940)

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the very large eye, the scape almost reaching the apex of the clypeus, the truncate antero-lateral pronotal projection, the glabrous pronotum, the long, subacute, brownish-black scutellar process and the hind femur being brown with two straw-colored annulations at the base and sub-apex. It most closely resembles M. modesta comb. nov., but is differentiated by the color pattern on the hind femur and the shape of the scutellar spine.

Other material
VESTITURE. Glabrous. Head with some fl at, curved 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 irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with short, curved setae on humeral angle and sparsely distributed along dorsal surface. Corium of hemelytron glabrous. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen with a few clubbed setae on each segment. HEAD. Elongate conical; maxillary plate rounded apically; scape not reaching apex of clypeus; eye distinctly projecting in dorsal view, less than 1/5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe level with postocular lobe.
LEGS. Same as genus description.

Female
Similar in size, shape and color to male except with wider abdomen.

Distribution
This species is only known from two localities in north Borneo, Sabah.

Remarks
Macrophysoderes histrionica comb. nov. shares the dark body color, head shape, general vestiture, and the shape of the parascutellar lobe with M. monticola comb. nov. and is thus transferred to the new genus to refl ect the hypothesized close relationship. Examination of the type series of M. histrionica comb. nov. showed that 3 of the paratypes match the holotype of M. modesta comb. nov. more closely than M. histrionica comb. nov. and these specimens are here treated under M. modesta comb. nov. (Miller, 1940)

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the elongate, conical head, the rounded maxillary plate, the anteocular lobe being almost level with the postocular lobe, the truncate antero-lateral rounded pronotal projections, the rounded parascutellar lobes that are skewed towards the median, the dark brown hind femur with a single basal straw-colored annulation, the apex of the scutellar process being straw-colored, the undulating connexivum, the cup-like sclerite with distinct medial process, and the fl ap-like prolongations of the phallosoma being plate-like and with a uniform basal margin. It is similar to M. monticola comb. nov. and M. histrionica comb. nov., but can be differentiated by the uniform color of the legs and connexiva, the margin of the pygophore posterior opening having a medial process and cup-like sclerite, and the shape of the plate-like prolongations of the phallosoma.
COLORATION (Figs 7,(10)(11). Dark brown. Head dark brown. Scape of antenna brown, straw brown basally, pedicel brown, basi-and distifl agellomere brown. Labium brown. Pronotum dark brown, scutellum dark brown, pleuron dark brown, sternum dark brown. Corium of hemelytron dark brown, membrane black to dark brown. Fore and mid legs dark brown, hind femur dark brown, hind tibia dark brown with straw-colored apex, hind tarsus and claw straw-colored. Abdomen dorsally orange-brown, ventrally light brown to brown with reddish-brown or dark brown lateral suffusion, connexivum dark brown with straw-colored posterior margin.
VESTITURE. Glabrous. Head with some fl at, curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, without pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with only short, sparse setae. Corium of hemelytron with short, sparse, adpressed setae. Legs with two rows of spines and setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum on abdomen with short, fi ne setae. HEAD. Elongate conical; maxillary plate rounded apically; scape not reaching apex of clypeus; eye distinctly projecting in dorsal view, less than 1/5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe level with postocular lobe.
THORAX. Antero-lateral paired projections truncate; surface of anterior lobe smooth, ridges almost obsolete; median pronotal depression contiguous with transverse sulcus; paramedian carina weakly defi ned; posterior lobe medially rugose; anterior pronotal lobe shorter than posterior lobe, narrower than posterior lobe, anterior lobe lower than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellar process long, with subacute apex; mesosternite with median irregular tuberculated protrusion between fore and mid coxae.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, with medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening with small medial process; cup-like sclerite with medial process; arms of basal plate rounded; ductifer membranous; endosomal struts conical, with subacute apex, basally divided into two arms; shape of dorsal phallothecal sclerite subacute, with lateral broad, plate-like prolongations.

Female
Similar to male.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is collected from both Sabah and Sarawak, Malaysia.

Remarks
This species is transferred to Macrophysoderes gen. nov. because of the similarities it shares with M. monticola comb. nov. and M. histrionica comb. nov. Examination of the type specimens of P. sibauana and P. trusana shows no morphological differences between them and they are hereby synonymized under M. modesta comb. nov. (Miller, 1940)

Diagnosis
This species is recognized among species of Macrophysoderes gen. nov. by the elongate conical head, the rounded maxillary plate, the anteocular lobe being almost level with the postocular lobe, the truncate antero-lateral pronotal projections, the rounded parascutellar lobes being skewed towards the median, the dark brown hind femur with a single basal straw-colored annulation, the apex of the scutellar process being straw-colored, the connexivum undulating, and the plate-like and angular fl ap-like prolongations of the phallosoma. It is similar to M. modesta comb. nov. and M. histrionica comb. nov., but can be differentiated by the color of the legs, scutellar spine and connexiva, the shape of the margin of the posterior opening of the pygophore and cup-like sclerite, and the shape of the plate-like prolongations of the phallosoma.
VESTITURE. Glabrous. Head with some fl at, curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, without pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with short, curved setae on humeral angle and sparsely distributed along dorsal surface. Corium of hemelytron with short, sparse, adpressed 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. HEAD. Elongate conical; maxillary plate rounded apically; scape not reaching apex of clypeus; eye distinctly projecting in dorsal view, less than 1/5 length of head, not reaching ventral margin of head in lateral view; height of anteocular lobe level with postocular lobe.
THORAX. Antero-lateral paired projections truncate; surface of anterior lobe smooth, ridges almost obsolete; median pronotal depression contiguous with transverse sulcus; paramedian carina weakly defi ned; posterior lobe medially rugose; anterior pronotal lobe shorter than posterior lobe, narrower than posterior lobe, anterior lobe lower than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellar process long, with subacute apex; mesosternite with median irregular tuberculated protrusion between fore and mid coxae.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded and rim ventrally sclerotized; arms of basal plate parallel to each other; ductifer membranous; endosomal struts conical, with subacute apex, basally divided into two arms; dorsal phallothecal sclerite subacute, with lateral broad, plate-like prolongations, broad plates angularly oriented towards basal plate.

Female
Similar to male in size, shape and color.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is distributed along the western area of peninsular Malaysia. Specimens from the state of Perak are larger than those from Selangor.

Remarks
Based on the phylogenetic analysis, this species forms a monophyletic clade with M. bengalensis comb. nov. as part of the Macrophysoderes gen. nov. clade. Other species closely resembling M. monticola comb. nov. are also transferred to Macrophysoderes gen. nov. based on the hypothesis that they are most closely related, namely M. modesta comb. nov. and M. histrionica comb. nov. Common characteristics shared among these three species include having an elongate conical head, truncate antero-lateral pronotal projections and a hemelytron with a dark brown membrane. Examination of the type specimens of P. dimidiata and P. luiana show no morphological differences except for size and are hereby synonymized with M. monticola comb. nov. This species and M. modesta comb. nov. are highly similar and diffi cult to distinguish other than based on the morphology of the male genitalia and on distribution.

Diagnosis
This genus is recognized among other genera of Physoderinae by the small size, the pair of prominent setigerous tubercles on the ventral surface of the postocular lobe, the lateral margins of the anterior pronotal lobe being almost straight and not curved, the semicircular shape of the scutellum, the scutellum having distinct paired lateral tubercles, the mesosternum having three longitudinal rows of setae and the armature of the fore femur consisting of two dense rows of setigerous tubercles. The genus is distinct from any other physoderine genus by the straight lateral margins of the anterior pronotal lobe, small size, uniform coloration, sparse setation over the entire body, paired projections on the scutellum, and the three longitudinal rows of setae on the mesosternum.

Etymology
The name combines Physoderes after the type genus of Physoderinae and the Latin noun "nanus", meaning dwarf, to indicate small body size. The gender is feminine.

Description
Female BODY LENGTH. Small, total length 7.66 mm.
VESTITURE. Sparsely setose. Head with some fl at, curved setae or with widespread short, fi ne, adpressed setae, ventral surface of postocular lobe with only a pair of prominent setigerous tubercles, with pair of long, straight setae wider apart on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with only short, sparse setae. Corium of hemelytron with short, sparse, adpressed setae. Legs with two rows of setigerous tubercles, tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen glabrous. HEAD. Short conical; maxillary plate apically truncate; scape not reaching apex of clypeus; eye hemispherical in dorsal view, about 1/ 5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe shorter than postocular lobe.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe with low ridges; median pronotal depression contiguous with transverse sulcus; paramedian carina weakly defi ned; posterior lobe medially weakly rugose; anterior pronotal lobe shorter than posterior lobe, narrower than posterior lobe, anterior lobe lower than posterior lobe in lateral view; paramedian lobe bell-shaped and skewed towards median; scutellum semicircular, scutellar process long, with subacute apex; mesosternite without median irregular tuberculated protrusion between fore and mid coxae.

Ecology
Unknown.

Distribution
Known only from the locality of the holotype of the type species, which was labelled as "New Guinea".

Remarks
Based on the phylogeny, N. dentiscutum comb. nov. is sister to all other physoderines except for the Neotropical and Afrotropical species. The isolated placement of N. dentiscutum comb. nov. in the phylogeny and its distinct morphology warrant the decision to erect a new genus to accommodate this species. This genus is described based on a female specimen as no males are known.

Diagnosis
As in generic description.

Diagnosis
This genus is recognized among other genera of Physoderinae by its small size (6.30-7.09 mm), brown to dark brown coloration, the scape surpassing the apex of the clypeus and being thicker than the remaining antennal segments, the obsolete antero-lateral pronotal projection, the rounded lateral margins of the anterior pronotal lobe, the anterior pronotal lobe being level with the posterior lobe, the acute scutellar spine, and the fore tibia bearing a few prominent processes with sharp, stout, setae. It is most similar to Physoderoides Miller (1955a), but is distinguished by the body size, head shape, the maxillary plate not being raised and the lack of antero-lateral pronotal projections.
VESTITURE. Sparsely setose with tuberculated, curved, short, setae. Surface of anterior pronotal lobe smooth or tuberculated. HEAD. Short conical; maxillary plate rounded apically; scape surpassing apex of clypeus; eye relatively large, hemispherical or projecting in dorsal view, not reaching ventral margin of head in lateral view; ocelli present; height of anteocular lobe level or shorter than postocular lobe.
THORAX. Antero-lateral paired projections obsolete; ridges of anterior pronotal lobe almost obsolete; median pronotal depression contiguous with transverse sulcus; paramedian carina weakly defi ned; length of anterior pronotal lobe variable, narrower than posterior lobe in both sexes, level with posterior lobe in lateral view; parascutellar lobe semicircular or quadrant-shaped, skewed towards median; scutellum triangular, scutellar process long, apex variable; mesosternite with median, irregular, tuberculated protrusion between fore and mid coxae.
ABDOMEN. Ovoid or elongate ovoid, with straight or rounded terminal margin; connexival margin smooth, posterior margin not elevated.
GENITALIA. Only examined for Pa. popeye sp. nov. (see species description below).

Female
Similar to males, but larger and with wider abdomen.

Ecology
No information is available for the Madagascan species, while Pa. popeye sp. nov. has been recorded on Pandanus (Pandanaceae).

Distribution
This genus has a disjunct distribution, with two species occurring in northeastern Madagascar and one species on the eastern edge of Papua New Guinea, on the Huon Peninsula.

Remarks
Based on the phylogenetic analysis, the new species Paraphysoderes popeye sp. nov. forms a monophyletic clade with Pa. peyrierasi and Pa. crassa. It shares the synapomorphies of the small body size and the fore tibia bearing a few prominent setigerous tubercles. Closer examination also revealed similarities in the shape of the head and pronotum, a similar acute scutellar process and the hemelytron not reaching the tip of the abdomen. This evidence led to classifying this new species as Paraphysoderes, even though this signifi cantly expands the distribution range of this genus, which was previously restricted to Madagascar.

Key to species of Paraphysoderes
1. Fore femur longer than mid femur, surpassing apex of clypeus, scutellar process acute and tapered, Cu and M veins not parallel, converging basally (Fig. 14

Diagnosis
This species is recognized among species of Paraphysoderes by its small size, brown to dark brown coloration, the elongate conical head, the scape surpassing the apex of the clypeus, the obsolete anterolateral pronotal projection, the anterior pronotal lobe being level with the posterior lobe, the scutellar spine being acute and tapered, the parascutellar spine being semicircular, the cubital (Cu) vein being angular at the junction of the connecting vein with the fi rst anal vein (1A), the distinctly incrassate fore femur, which is longer than the mid femur, the fore tibia bearing a few prominent processes with sharp, stout, setae, the mid femur with prominent processes with peg-like setae on the anterior surface, and the abdominal spiracles turreted and being located directly ventral to the connexival margin.

Etymology
The name popeye is a noun in apposition to describe the uniquely enlarged fore-arms of this species that is similar to the fi ctional cartoon character Popeye the Sailor Man. COLORATION (Fig. 14). Brown to dark brown. Head brown or dark brown. Scape and pedicel of antenna light brown with brown suffusion, disti-and basifl agellomeres straw-colored. First segment of labium brown, second segment basally brown, medially straw-colored, apically brown, third segment brown. Thorax dark brown. Corium of hemelytron dark brown, membrane dark brown. Fore and mid femora and tibiae of legs light brown to brown, tarsi and claws light brown, hind femur dark brown with medial light brown annulation, hind tibia light brown to dark brown, hind tarsus and claw light brown. Abdomen dorsally reddish-orange, ventrally light brown with brown suffusion, connexivum brown with a lighter margin, exposed part of pygophore dark brown.

Holotype
VESTITURE. Sparsely setose. Head with widespread curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, without pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, posterior lobe with short, curved, setigerous tubercles on humeral angle and glabrous on dorsal surface. Corium of hemelytron with short, sparse, adpressed setae. Legs with two rows of spines and setigerous tubercles, tibia with a few prominent tuberculated, stout, sharp setae. Connexival margin of abdomen sparsely setose, without prominent setae.
HEAD. Elongate conical; maxillary plate rounded apically; scape surpassing apex of clypeus; eye hemispherical in dorsal view, about 1/ 5 length of head, not attaining ventral margin of head in lateral view; height of anteocular lobe level with postocular lobe.
HEMELYTRON. Not reaching tip of abdomen.
GENITALIA. Anterior margin of mediosternite 8 sharply emarginate, without medial apodeme; transverse bridge of pygophore broad, margin of anterior opening rounded, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded, with sclerotized paired latero-ventral slight protuberances; arms of basal plate parallel to each other; ductifer with sclerotized rounded ring; endosomal struts tube-like, with medial sclerotization and with a T-shaped apex; shape of dorsal phallothecal sclerite subacute with narrow, lateral, band-like prolongations, sclerotized band extended horizontally towards ventral surface of phallosoma.

Female
Similar in morphology to males, but much larger (7.89 mm average total length) with wider abdomen (2.56 mm for males, 3.32 mm average width for females). Because one such female was collected in the same collecting event as the holotype male, these females are here treated as the same species.

Ecology
This species has been recorded on Pandanus (Pandanaceae).

Distribution
This species is distributed in the eastern edge of Papua New Guinea around the Huon Peninsula.

Remarks
A damaged female specimen from Fiji (not described here), that is most likely not the same species as Pa. popeye sp. nov. but likely belongs to Paraphysoderes, suggests an even more extended and disjunct distribution of this genus and potential undiscovered diversity in the Pacifi c region. Note: The current color of the holotype has been altered and is different from the image provided here due to treatment with KOH solution for dissection of the male genitalia.

Diagnosis
This genus is recognized among other genera of Physoderinae by the scape reaching the apex of the clypeus, the parascutellar lobe being rounded and skewed towards the median, the males having an enlarged anterior pronotal lobe, wider than or equal to the width of the posterior lobe (except in P. nigripennis sp. nov., P. ractepilosa sp. nov. and P. muluensis sp. nov.), the median pronotal depression not being contiguous with the transverse sulcus (males) and by males having a membranous ductifer. This genus is most similar to Epiroderoides Villiers, 1962, but can be differentiated from it based on the anterior pronotal shape in males and the shape of the parascutellar lobes.
VESTITURE. Sparsely to densely setose with curved, fi ne or erect setae, with pair of long, straight setae on posterior margin of postocular lobe (except in P. nigripennis sp. nov. and P. nigroalbus).
HEAD. Elongate or short, conical; maxillary plate most often rounded apically; scape never distinctly surpassing apex of clypeus; eye hemispherical in dorsal view except in P. nigripennis sp. nov. and P. tricolor sp. nov., less than 1/ 5 length of head and never reaching ventral margin of head in lateral view; ocelli present; height of anteocular lobe shorter than postocular lobe except in P. anamalaiensis sp. nov. and P. mysorensis sp. nov.
THORAX. Shape of antero-lateral paired pronotal projection variable; surface of anterior pronotal lobe most often with low ridges; median pronotal depression not contiguous with transverse sulcus except in species without enlarged anterior pronotum; paramedian carina variable; posterior lobe medially rugose; length of anterior pronotal lobe variable, wider and higher than or equal to posterior lobe except in P. nigripennis sp. nov., P. ractepilosa sp. nov. and P. muluensis sp. nov.; parascutellar lobe rounded and sometimes skewed towards median, except in P. brevipennis sp. nov.; scutellum rounded triangular except in P. azrael and P. curculionis, length of scutellar process and shape of apex variable; mesosternite with median irregular tuberculated protrusion between fore and mid coxae.
ABDOMEN. Elongate ovoid except in P. brevipennis sp. nov. and P. curculionis, with rounded terminal margin except in P. ractepilosa sp. nov.; shape of connexival margin variable, posterior margin only slightly elevated in P. nigripennis sp. nov.
GENITALIA. Variable (see species descriptions for details).

Female
Females with anterior pronotal lobe narrower and lower than posterior lobe, with median pronotal depression contiguous with transverse depression.

Ecology
Species from this genus are found in a variety of living and dead plant materials and in the undergrowth. Specimens have been collected from pitfall traps and carrion traps and have been found at up to 1200 m elevation.

Distribution
This genus is the most widespread of the subfamily, and species distributions cover the entire known range of Physoderinae in the Old World except in the Afrotropics. Species can be found in Madagascar, the Indian subcontinent, peninsular Malaysia, Sumatra, Java, Borneo, the Philippine archipelago, Sulawesi, Papua New Guinea, the Solomon Islands and the Fiji Islands.

Diagnosis
This species is recognized among species of Physoderes by the scape reaching the apex of the clypeus, the small eye, the acute and anteriorly oriented antero-lateral projection of the anterior pronotal lobe, the wide anterior pronotal lobe, which is equal to or wider than the posterior lobe in males and only slightly narrower than the posterior lobe in females, the rounded parascutellar lobe, the dark brown scutellum with the apex thickened and raised, the straw-colored, rounded and short scutellar process, and the hemelytron reaching the tip of the abdomen. This species is most similar to P. mysorensis sp. nov., but is differentiated by the smaller size (8.91-9.82 mm), having no apodeme on mediosclerite 8, the narrow transverse bridge of the pygophore, and the margin of the anterior opening of the pygophore being angular in lateral view and with a membranous ductifer.

Etymology
The name anamalaiensis is a noun in apposition as a tribute to the type locality Anamalai Hills, Tamil Nadu, India. COLORATION (Figs 14, 17). Dark brown. Head dark brown. Scape of antenna brown, pedicel brown basally, straw-colored apically, basifl agellomere brown, distifl agellomere basally brown, apically straw-colored. First segment of labium brown, second segment basally brown, medially straw-colored, apically brown, third segment brown. Pronotum of thorax dark brown, scutellum dark brown, scutellar process strawcolored, pleuron dark brown, sternum dark brown. Corium of hemelytron dark brown, membrane dark brown. Legs dark brown. Abdomen dorsally yellowish-orange, ventrally dark brown with suffusion of straw-color and brown, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore dark brown.

Holotype
VESTITURE. Sparsely setose. Head with widespread curved 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 irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, 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, tibiae with regular rows of tuberculated, stout, sharp setae. Connexival margin of abdomen with no prominent setae. HEAD. Elongate conical; maxillary plate rounded apically; scape reaching apex of clypeus; eye hemispherical in dorsal view, less than 1/ 5 length of head; height of anteocular lobe level with postocular lobe.
THORAX. Antero-lateral paired projections acute; surface of anterior lobe with low ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defi ned; anterior pronotal lobe longer than posterior lobe, wider than or equal in width to posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded; scutellar process short, apex rounded.

LEGS. Same as genus description.
ABDOMEN. Connexival margin slightly undulating, posterior margin not elevated.

Female
Similar to males except with anterior pronotal lobe slightly narrower than posterior lobe and median pronotal depression contiguous with transverse sulcus.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is known only from the area around the type locality in the Anamalai Hills, Madras, Tamil Nadu, India.

Remarks
This species possesses the enlarged anterior pronotal lobe characteristic of Physoderes along with the associated pronotal modifi cations, and is thus placed in this genus. The females possess an enlarged anterior pronotal lobe, although not as exaggerated as that of the males, which is quite unique. The species is most likely closely related to the other Physoderes species native to India, P. mysorensis sp. nov., described below. Kirkaldy, 1905 Figs 8-9, 14, 17, 20;Appendix Physoderus [sic] azrael Kirkaldy, 1905: 339.

Diagnosis
This species is recognized among other species of Physoderes by the head being anteriorly light brown and posteriorly brown, the pronotum dark brown straw-colored or with a light brown humeral tubercle, the posterior margin, parascutellar lobe, scutellar process and base of the corium light brown, 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 parascutellar lobe being rounded and skewed towards the median, the semicircular scutellum, the spatulate and apically rounded scutellar process, the pronotum fl attened in lateral view and the hemelytron reaching the tip of the abdomen. In the male genitalia, the following features are also diagnostic: the extremely narrow transverse bridge of the pygophore and the apically truncate endosomal struts, which are basally divided into two arms converging medially to form basally a ridge. This species is most similar to P. curculionis but can be differentiated by the coloration, elevation of the pronotum, shape of the abdomen and shape of the endosomal struts in males.
VESTITURE. Densely setose. Head with some curved setae and short, fi ne, adpressed setae, ventral surface of postocular lobe with two rows of small, setigerous tubercles, with pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with short, sparse setae intermixed with short, tuberculated, curved setae. Corium of hemelytron with short, sparse, adpressed 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 defi ned; anterior pronotal lobe equal in length to posterior lobe, equal in 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.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore extremely narrow, only consisting of margin of anterior opening of pygophore, margin of anterior opening angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded, with adjacent paired round protuberances; arms of basal plate converging; ductifer with sclerotized, rounded ring; endosomal struts apically truncate, basally divided into two arms converging medially to form a tapered ridge basally; shape of dorsal phallothecal sclerite subacute with lateral broad, plate-like prolongations.

Ecology
This species has been collected most commonly on or inside Heliconia fl owers (Heliconiaceae), but also on Pandanus (Pandanaceae) in forests, on foliage of Avocado pear (Lauraceae), in rotting tissue of the trunk of Roystonea regia (Arecaceae), on Nypa fruticans (Arecaceae), on Cocos nucifera (Arecaceae), under sacking bands on trees, logs near the coast, in coastal litter, under dead bark and on Musa (Musaceae).

Distribution
This species is mostly found on New Britain Island and the Solomon Islands archipelago, but is also found on Fiji Island, Papua New Guinea, Sabah (Borneo) and Pulau Larat (Indonesia). This species has the eastern-most distribution of all physoderines with specimens found in Fiji.

Remarks
This species is placed in the Physoderes clade based on the phylogenetic analysis above. It is most closely related to P. curculionis, with which it shares the semicircular scutellum and the very narrow transverse bridge on the pygophore. 14,17,[19][20]; Appendix

Diagnosis
This species is recognized among other species of Physoderes by the small size, the head and pronotum being densely covered with long, curved setae, the elongate conical head, the scape reaching the apex of the clypeus, the enlarged anterior pronotal lobe, the wide posterior pronotal lobe (males), the posterior pronotal lobe and parascutellar lobe being reduced in size, the brachypterous wing type and the ovoid abdomen. Among male genitalic features, the following are also diagnostic: the narrow transverse bridge of the pygophore, the endosomal struts being apically bulbous and basally divided into two plates converging to form a ridge medially, and the dorsal phallothecal sclerite shaped like a thin moustache. This species is distinct by being the only brachypterous Physoderes species and it is restricted to Guadalcanal of the Solomon Islands archipelago.

Etymology
The name brevipennis is chosen after the Latin adjective "brevipennis", meaning short-winged, to describe the short wings of this species.
VESTITURE. Densely setose. Head with widespread curved 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. HEAD. Elongate conical; maxillary plate rounded apically; scape just surpassing 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 strongly defi ned; anterior pronotal lobe longer and wider than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe bell-shaped, skewed towards median; scutellum rounded triangular, scutellar process short, apex subacute.
HEMELYTRON. Not reaching tip of abdomen.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore narrow, margin of anterior opening angular, apodeme present, 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 bulbous, basally divided into two plates converging to form a ridge medially; shape of dorsal phallothecal sclerite shaped like a thin moustache.

Female
Similar to males, with abdomen wider (average width 3.80 mm).

Ecology
This species has most often been collected using pitfall and carrion traps and one specimen was found on rotting pawpaw (Caricaceae).

Distribution
This species is restricted to Guadalcanal of the Solomon Islands archipelago.

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.
COLORATION (Figs 14, 16-17). Brown. Head brown. Scape, pedicel and basifl agellomere of antenna light brown, distifl agellomere 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, fi ne, 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 defi ned; 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.
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 confi rmed 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. Cao et al., 2011 Figs 13-14, 17 Physoderes esakii Cao, Tomokuni & Cai, 2011: 23, fi gs 1-12. Epirodera latithorax Esaki, 1931: 212 (nomen nudum).

Re-diagnosis
This species is recognized among species of Physoderes by its short hemelytra, not reaching the abdominal apex, the slender scutellar spine, the round parascutellar lobe and the simple phallosoma without lateral prolongations as illustrated in the original description and color images provided by Cao et al. (2011). This species is most similar to P. fuliginosa but can be differentiated on these characters. Cao et al. (2011) for full description.

Distribution
This species is only recorded from Botel Tobago (Orchid Island [Lanyu], Taiwan).

Remarks
A re-diagnosis is provided here, as the original diagnosis (under Remarks in Cao et al. 2011) refers to the sexually dimorphic enlargement of the anterior pronotal lobe also present in other species of Physoderes. Two female specimens likely to be conspecifi cs collected by J.F. Tsai on Botel Tobago island share the same short hemelytron not reaching the abdominal apex, but without an enlarged anterior pronotal lobe and with a rounder abdomen (J.F. Tsai and D. Rédei, pers. comm.; images examined here). (Stål, 1870) Figs 8, 13-14, 17, 19; Appendix Epirodera fuliginosa Stål, 1870: 693.

Diagnosis
This species is recognized among species of Physoderes by the elongate head, the scape reaching the apex of the clypeus, the hemispherical eye that is not distinctly protruding, the acute and diverging anterolateral projection of the anterior pronotal lobe, the anterior pronotal lobe being wider than the posterior lobe, the rounded parascutellar lobe, the slender and straw-colored scutellar process, the pygophore apical margin of the posterior opening having a short medial process and the phallosoma lacking fl aplike prolongations. This species is most similar to P. minime sp. nov., but can be differentiated from it by the larger size, darker coloration, the hemelytron close to or attaining the tip of the abdomen, the presence of an apical median process on the posterior opening of the pygophore, and the lack of fl ap-like prolongations of the phallosoma.

Material examined
Holotype PHILIPPINES: ♀, specimen image made available by T. Ishikawa, label information not examined here (NHRS).
COLORATION (Figs 14, 17). Dark brown. Head dark brown. Scape and pedicel of antenna light brown and brown, basifl agellomere brown, distifl agellomere basally brown, apically straw-colored. First segment of labium brown, second segment basally and apically brown, medially straw-colored, third segment brown. Pronotum of thorax dark brown with light brown markings, scutellum basally dark brown with scutellar process straw-colored, pleuron straw-colored with brown and dark brown patterns, sternum dark brown with brown suffusion. Corium of hemelytron brown to dark brown, membrane dark brown. Femur and tibia of legs straw-colored with medial and apical brown annulations, tarsus and claw light brown. Abdomen dorsally yellowish-orange, ventrally straw-colored with sub-lateral dark brown patterns, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore brown.
VESTITURE. Sparsely setose. Head with widespread curved 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 irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, 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 no prominent setae. HEAD. Elongate 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 defi ned; anterior pronotal lobe longer and wider than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded; scutellum rounded triangular, scutellar process long, apex subacute.
LEGS. Same as genus description.

Female
Similar to male, except with anterior pronotal lobe (2.39 mm average) distinctly narrower than posterior lobe (3.20 mm average) and with median pronotal depression contiguous with transverse sulcus.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is widespread across the Philippines island archipelago and also recorded from Borneo, parts of the Indonesian archipelago, Papua New Guinea and the Solomon Islands.

Remarks
The holotype of P. fuliginosa is a female with a narrow anterior pronotal lobe and unknown locality, which makes it diffi cult to associate it with a male. Multiple male specimens found on Mindanao with Fig. 17. Ventral habitus images for species of Physoderes Westwood, 1845 (partial). Scale bars = 2 mm. corresponding females that match the holotype morphology confi rm that the holotype is most likely from the Philippines region and the redescription here is based on the males from Mindanao. The description of P. fuliginosa is diffi cult, as specimens from the Philippines show variation in overall size and coloration, but do not differ morphologically. Dissections of male genitalia from different geographic localities also do not show any variation. Hence, P. fuliginosa is here considered to be highly variable in size and general coloration.

Diagnosis
This species is recognized among species of Physoderes by the elongate head, which is as long as the pronotum, the scape not reaching the apex of the clypeus, the hemispherical eye being slightly fl attened, the antero-lateral projection of the anterior pronotal lobe subacute and diverging, the anterior pronotal lobe being wider than the posterior lobe, the rounded parascutellar lobe, the scutellar process being slender and straw-colored and the short hemelytron which does not reach the tip of the abdomen. This species is most similar to P. fuliginosa, but can be differentiated from it by the smaller size, yellowish coloration, the short hemelytron and by having a smooth apical margin on the posterior opening of the pygophore, as well as a small apical dorsal phallothecal sclerite.

Etymology
The name minime is a noun in apposition, after the fi ctional movie character Mini-Me from the Austin Powers movie series. The name given is to describe the miniaturized resemblance of this species to the co-occurring larger P. fuliginosa. COLORATION (Figs 14, 17). Yellowish-brown. Head yellowish to light brown. Scape, pedicel and distifl agellomere of antenna straw-colored, basifl agellomere basally brown, apically straw-colored. First segment of labium light brown, second segment straw-colored, third segment brown. Pronotum of thorax brown with straw-colored markings along ridges, scutellum basally brown, apically strawcolored including scutellar process, pleuron straw-colored with brown suffusion, sternum brown. Corium of hemelytron light brown to brown, membrane light brown to brown. Femora and tibiae of legs straw-colored with sub-basal and apical light brown annulations, tarsi and claws straw-colored. Abdomen dorsally yellowish orange, ventrally straw-colored with lateral brown suffusion, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore straw-colored brown.

Holotype
VESTITURE. Sparsely setose. Head with widespread curved 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 irregular row of tuberculated, short, curved setae on lateral margins and fi ne, adpressed setae on dorsal surface, posterior lobe with short, curved, setigerous tubercles on humeral angle and glabrous on 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 no prominent setae. HEAD. Elongate conical; maxillary plate rounded apically; scape not reaching apex of clypeus; eye hemispherical in dorsal view.
THORAX. Antero-lateral paired projections subacute; surface of anterior lobe with low ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defi ned; anterior pronotal lobe longer and wider than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded; scutellum rounded triangular, scutellar process long, apex subacute.
HEMELYTRON. Not attaining tip of abdomen.
LEGS. Same as genus description.
GENITALIA. Transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded with sclerotized paired latero-ventral slight protuberance; arms of basal plate converging; ductifer membranous; endosomal struts apically spatulate, basally divided into two plates; shape of dorsal phallothecal sclerite subacute with short, apical lobes and rounded lateral plates at apex.

Female
Similar to males except anterior pronotal lobe (2.07 mm average) narrower than posterior lobe (2.60 mm average) and median pronotal depression contiguous with transverse sulcus.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is recorded from Luzon Island and Panay Island of the Philippines archipelago. Most specimens were collected from two localities: Mt Banahao and Mt Makiling in Luzon.

Remarks
This species differs from the co-occurring P. fuliginosa based on the diagnostic characters listed above and is placed within Physoderes based on the phylogenetic analysis conducted above.

Diagnosis
This species is recognized among congeners by the males having the anterior pronotal lobe narrower than the posterior lobe, the median pronotal depression being contiguous with the transverse sulcus, the fore femur slightly incrassate and by the narrow transverse bridge of the pygophore. This species is distinguished from other species of Physoderes by the distinctly small size and the anterior pronotal lobe not being infl ated in males.

Etymology
The name muluensis is a noun in apposition derived from the type locality Gunung Mulu National Park, Sarawak. COLORATION (Fig. 15). Brown. Head brown, lighter towards apex. Scape of antenna brown, pedicel brown with apex straw brown, basi-and distifl agellomeres basally brown and apically straw brown or straw brown entirely. First segment of labium brown, second segment gradation from brown to straw-colored, third segment straw-colored. Pronotum of thorax brown, sometimes posterior pronotal lobe lighter brown, scutellum brown, pleuron brown, sternum brown. Corium of hemelytron brown, membrane brown. Fore femur of legs brown entirely or basally dark brown, fore tibia straw-colored, basally brown, tarsus and claws straw-colored, mid femur entirely brown or brown with basally straw brown coloration, mid tibia, tarsus and claws brown and straw-colored, hind femur brown and basally straw-colored or straw-colored with medial brown band, hind tibia brown and straw brown, tarsus and claws straw brown. Abdomen dorsally orange-yellow, ventrally brown, connexivum brown with posterior third light brown, exposed part of pygophore brown.

Holotype
VESTITURE. Sparsely setose. Head with some fl at, curved setae or with widespread short, fi ne, 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 irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, posterior lobe with only short, sparse setae. Corium of hemelytron with short, sparse, adpressed setae. Legs with two rows of spines and setigerous tubercles, tibia with a few prominent tuberculated, stout, sharp setae. Posterior margin of connexivum of abdomen with long, fi ne setae.
HEAD. Elongate conical; maxillary plate truncate 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 contiguous with transverse sulcus; paramedian carina weakly defi ned; anterior pronotal lobe of equal length to and narrower than posterior lobe, level with posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum rounded triangular, scutellar process short, apex rounded or subacute.
HEMELYTRON. Not attaining tip of abdomen.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening with slight medial protuberance; cup-like sclerite apically rounded, with adjacent paired round protuberances; arms of basal plate parallel to each other; ductifer membranous; endosomal struts conical, subacute apex, divided into two arms basally; shape of dorsal phallothecal sclerite subacute with lateral broad, plate-like prolongations, broad plates angularly oriented towards basal plate.
COLORATION (Figs 15, 18). Dark brown. Head dark brown. Scape, pedicel and basifl agellomere of antenna brown, distifl agellomere basally brown, apically straw-colored. First segment of labium basally brown, apically straw-colored, second segment straw-colored, third segment brown. Pronotum of thorax dark brown, parascutellar lobes lighter in color, scutellum dark brown, scutellar process straw-colored, pleuron dark brown with brown and straw-colored suffusion, sternum dark brown. Corium of hemelytron dark brown, membrane dark brown. Femur of legs straw-colored with medial and apical brown annulations, tibia brown with basal and apical straw-colored brown annulations, tarsus and claw straw-colored. Abdomen dorsally yellowish-orange, ventrally dark brown with suffusion of straw-color and brown, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore brown.
VESTITURE. Sparsely setose. Head with widespread curved setae, ventral surface of postocular lobe with two rows of small, setigerous tubercles, with pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, 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 no prominent setae. HEAD. Elongate conical; maxillary plate rounded apically; scape reaching apex of clypeus; eye hemispherical in dorsal view, less than 1/ 5 length of head; height of anteocular lobe level with postocular lobe.
THORAX. Antero-lateral paired projections acute; surface of anterior lobe with low ridges; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defi ned; anterior pronotal lobe longer and wider than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded; scutellum rounded triangular, scutellar process short, apex subacute; mesosternite with median irregular tuberculated protrusion between fore and mid coxae.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, with medial apodeme; transverse bridge of pygophore broad, margin of anterior opening rounded, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded and rim ventrally sclerotized; arms of basal plate converging; ductifer with sclerotized angular ring; endosomal struts conical, with subacute apex, divided into two arms basally; shape of dorsal phallothecal sclerite subacute with lateral broad, platelike prolongations, with short, rounded lateral plates.

Female
Similar to males except anterior pronotal lobe slightly narrower than posterior lobe and median pronotal depression contiguous with transverse sulcus.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is known from two localities in Southwestern India: Agumbe Ghat, Mysore District, Karnataka and Ponmudi Range, Trivandrum District, Kerala.

Remarks
This species possesses the enlarged anterior pronotal lobe characteristic of Physoderes, along with the associated pronotal modifi cations, and is thus placed in this genus. The females possess an enlarged anterior pronotal lobe, although not as exaggerated as that of the males, which is quite unique. It is most likely to be closely related to the other Physoderes species native to India, P. anamalaiensis sp. nov. described above.

Diagnosis
This species is recognized among congeners by the elongate conical head, which is densely covered in fi ne, curved setae, the scape reaching the apex of the clypeus, the eye being distinctly projecting, the antero-lateral pronotal projection being truncate, the anterior pronotal lobe distinctly narrower than the posterior lobe and covered with fi ne, curved setae along its ridges, the parascutellar lobe rounded and skewed towards median, the apex of the scutellar process straw-colored, the costal margin of the hemelytron black, the connexivum undulating with the posterior margin slightly elevated and beset with short, curved setae, the arms of the basal plate of the articulatory apparatus of the male genitalia being curved and the dorsal phallothecal sclerite being subacute, with lateral rounded plates extending to the lateral surface.

Etymology
The name nigripennis is after the Latin adjective "nigripennis", meaning with black wings or feathers, to describe the overall black coloration of the hemelytron.
COLORATION (Figs 15,18). Brown and dark brown. Head brown with straw-colored setae. Antenna light brown. First segment of labium brown, second segment straw-colored, apically brown, third segment brown. Anterior lobe of pronotum dark brown with brown or light brown markings, posterior lobe brown, scutellum basally brown, apically straw-colored including scutellar process, pleuron brown, sternum brown. Corium of hemelytron reddish-brown with black costal margin, membrane brownishblack. Femora of legs straw-colored with medial and apical brown annulations, tibiae basally brown, apically straw-colored, tarsi and claws light brown. Abdomen dorsally yellowish-orange, ventrally light brown medially and straw-colored laterally with brown suffusion, anterior two-thirds of connexivum brownish-black, posterior third straw-colored, exposed part of pygophore dark brown.
VESTITURE. Generally setose. Head with widespread curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, without pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and curved setae on anterior portion of dorsal surface, posterior lobe with widespread short, curved setae. 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. Posterior margin of connexivum of abdomen with short, curved setae. HEAD. Elongate conical; maxillary plate truncate apically; scape reaching apex of clypeus; eye distinctly projecting in dorsal view, less than 1/ 5 length of head.
THORAX. Antero-lateral paired projections truncate; surface of anterior lobe with raised ridges; median pronotal depression contiguous with transverse sulcus; paramedian carina strongly defi ned; anterior pronotal lobe shorter and narrower than posterior lobe, lower than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum rounded triangular, scutellar process long, apex subacute.
LEGS. Same as genus description.
GENITALIA. Transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded, with adjacent paired round protuberances; arms of basal plate rounded; ductifer membranous; endosomal struts conical, with subacute apex, divided into two arms basally; shape of dorsal phallothecal sclerite subacute with lateral broad, plate-like prolongations, with short, rounded lateral plates at apex.

Female
Similar to male.

Ecology
This species was collected at relatively high altitudes of 920-1200 m.

Distribution
This species is found only in the central west region of Sumatra Island, where most specimens were collected from Bukit Tinggi, formerly Fort de Kock.

Remarks
This species is the most commonly collected on the island of Sumatra. It is placed within the Physoderes clade in the phylogenetic analysis above because of the shared synapomorphies of having the scape reaching the apex of the clypeus, rounded parascutellar lobes skewed toward the median and a membranous ductifer of the male genitalia. Fig. 19. Dorsal view of the phallus in species of Physoderes Westwood, 1845 (partial). Breddin, 1903 Figs 15, 20;Appendix Physoderes nigroalbus Breddin, 1903: 126.

Diagnosis
This species is recognized among species of Physoderes by the short and conical head, the scape not reaching the apex of the clypeus, the eye hemispherical and slightly fl attened, the submedian pronotal carina strongly defi ned, the scutellum and scutellar process brown, the corium dark brown with the membranous portion basally dark brown including the external cell, and areas directly adjacent to the medial vein apical extension and fi rst anal vein apex with remaining membrane off-white. This species is most similar to P. tricolor sp. nov., but can be differentiated from it by its size, coloration of the head, pronotum and scutellar process, and the shape of the eye.

Redescription
Female BODY LENGTH. Medium, total length 9.74 mm. COLORATION (Fig. 15). Brown. Head brown. Scape and pedicel of antenna light brown to brown, basifl agellomere brown, distifl agellomere basally brown, apically straw-colored. First segment of labium light brown, second segment straw-colored, apically brown, third segment brown. Anterior lobe of pronotum dark brown, posterior lobe brown, scutellum brown with dark brown scutellar process, pleuron brown with straw-colored suffusion, sternum brown to dark brown. Corium of hemelytron dark brown, membrane basally dark brown, apically off-white. Femur and tibia of legs straw-colored with basal, medial and apical brown annulations, tarsus and claw light brown. Abdomen dorsally yellowishorange, basally straw-colored with brown and dark brown patterns, anterior half of connexivum dark brown, posterior half straw-colored.
VESTITURE. Sparsely setose. Head with widespread curved setae, ventral surface of postocular lobe with sparse, setigerous tubercles, without pair of long, straight setae on postocular lobe posterior to ocelli. Anterior lobe of thorax with irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, 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 no prominent setae. HEAD. Short conical; maxillary plate rounded apically; scape not 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 contiguous with transverse sulcus; paramedian carina strongly defi ned; anterior pronotal lobe shorter and narrower than posterior lobe, lower than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum rounded triangular, scutellar process long, apex subacute.
LEGS. Same as genus description.

Male
Unknown.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is only known from the type locality in northeast Sumatra, Indonesia.

Remarks
This specimen was found by Stephan Blank at the DEI without a type label and labeled as Epirodera palliderostris [sic]. It is here interpreted as the type specimen or part of a syntype series that Breddin used to describe P. nigroalbus. Because Breddin did not label his types nor mention where they were deposited, this is currently the best assumption. Further evidence is given by the exact match in the locality information provided in Breddin's description and that on the specimen label, the lack of a range of measurements to suggest he examined more than one specimen, and matching description of the black and white hemelytron, measurement and sex. This specimen is thus designated as the lectotype for P. nigroalbus. No matching male specimens have been located and the redescription here is based on the single female lectotype specimen. Westwood, 1845 Figs 8, 13, 15-16, 19

Diagnosis
This species is recognized among congeners by the short head, the scape shorter than the apex of the clypeus, the parascutellar lobe being rounded and skewed towards the median, the anterior pronotal lobe wider than the posterior lobe (males only), the median pronotal depression not contiguous with the transverse sulcus (males), the hemelytron reaching the tip of the abdomen, the hind wings being bright yellow and sometimes apically brown, the cup-like sclerite with rounded apex and adjacent paired protuberances, and the plate-like prolongations of the phallosoma with subacute apex and sharp lateral extensions. This species is similar to P. tricolor sp. nov. but can be differentiated from it based on the head and pronotal color patterns, hemelytron color, cup-like sclerite shape and the shape of the prolongations of the phallosoma.
of labium light brown to brown, second segment straw-colored, third segment brown. Pronotum of thorax dark brown with straw-colored markings along ridges, scutellum basally dark brown, apically straw-colored, pleuron straw-colored with brown and dark brown patterns, sternum dark brown. Corium of hemelytron light brown to brown, membrane dark brown. Femur of legs straw-colored with subbasal and apical brown annulations, tibia brown with straw-colored base, tarsus and claw straw-colored. Abdomen dorsally yellow, ventrally straw-colored with sub-lateral dark brown patterns, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore straw-colored.
VESTITURE. Sparsely setose. Head with some curved setae and short, fi ne, 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 irregular row of tuberculated, short, curved setae on lateral margins and short, setigerous tubercles dispersed on dorsal surface, posterior lobe with only short, sparse setae. 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 no prominent setae.
HEAD. Elongate conical; maxillary plate rounded apically; scape not 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 defi ned; anterior pronotal lobe equal in length to and wider than posterior lobe, higher than posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum rounded triangular, scutellar process long, apex subacute.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 sharply emarginate, without medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening smooth; cup-like sclerite apically rounded with adjacent paired round protuberances; arms of basal plate parallel to each other; ductifer membranous; endosomal struts with tapered apex and divided into two arms basally; shape of dorsal phallothecal sclerite subacute with lateral broad, platelike prolongations, broad plate with a subacute apex and sharp lateral extensions.

Female
Females have a narrower anterior pronotal lobe with median pronotal depression contiguous with transverse sulcus and anterior lobe lower than the posterior lobe.

Ecology
Not much is recorded about the biology of this species. One specimen was collected from under sheaths of bamboo (Poaceae), another one was collected from multi-storey evergreen forest using 'canopy trap fi sh'. This species has been collected from a range of mid-level elevations from 227 to 1496 m.

Distribution
This species is relatively widespread and can be found on peninsular Malaysia and the islands of Sumatra, Java and Sulawesi.

Remarks
Examination of the holotype of P. fl avipennis and the syntypes of P. brunneus indicate that both represent females of P. notata that have the narrower anterior pronotal lobe. Similarly, the holotype of P. corporaali shares the same diagnostic characters and color patterns on the head and pronotum with P. notata. Although the holotype of P. buruensis has a darker pronotum, all other characters are consistent with those of P. notata. The difference in pronotal coloration is hence regarded here as a color variant of the species, possibly unique to the Buru island population. All four species are hereby synonymized under P. notata.

Diagnosis
This species is recognized among species of Physoderes by the body being covered with long, erect setae, the head being elongate conical, the scape not reaching the apex of the clypeus, the anterior pronotal lobe being narrower than the posterior lobe, the anterior pronotal lobe with raised ridges and strongly defi ned submedian carina, the parascutellar lobe rounded, the scutellar process slender, the fore femur slender and only slightly incrassate, the short hemelytron not reaching the tip of the abdomen, the connexival margin undulating and the females with a wider connexivum than in males. Additional diagnostic features are found on the male genitalia: the cup-like sclerite with rounded apex and broad adjacent sclerotized rounded protuberances, the apical margin of the posterior opening of the pygophore with medial process, the arms of the basal plate curved to form a rounded foramen and the apex of the dorsal phallothecal sclerite subacute, with broad lateral prolongations that project angularly towards the apex of the phallosoma. This species most closely resembles P. fuliginosa, but can be differentiated from it by the long, erect setae, the shape of the pronotum, parascutellar lobe and connexivum, as well as the hemelytron length.

Etymology
The name ractepilosa is a noun in apposition to describe the erect setation throughout the body of this species.
VESTITURE. Generally densely setose. Head with widespread long, erect 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 irregular row of tuberculated, long, erect setae on lateral margins and on dorsal ridges, posterior lobe with long, erect setae on humeral angle and sparsely distributed along dorsal surface. Corium of hemelytron with long, erect setae. Legs with two rows of spines and setigerous tubercles, tibia with regular rows of tuberculated, stout, sharp setae. Posterior margin of connexivum of abdomen with long, fi ne setae.
HEAD. Elongate conical; maxillary plate truncate apically; scape not 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 raised ridges; median pronotal depression contiguous with transverse sulcus; paramedian carina strongly defi ned; anterior pronotal lobe shorter and narrower than posterior lobe, lower than posterior lobe in lateral view; parascutellar lobe rounded; scutellum rounded triangular, scutellar process long, apex subacute.
HEMELYTRON. Not reaching tip of abdomen.
LEGS. Same as genus description.

Male
Similar to females, but do not seem to have as many long, erect setae and connexivum not as wide.
GENITALIA. Anterior margin of mediosternite 8 sharply emarginate, with medial apodeme; transverse bridge of pygophore broad, margin of anterior opening angular, apodeme present, apical margin of posterior opening with small medial process; cup-like sclerite apically rounded, with adjacent paired round protuberances; arms of basal plate rounded; ductifer membranous; apex of endosomal struts tapered and divided into two arms basally; shape of apex of dorsal phallothecal sclerite subacute with broad lateral prolongations that project angularly towards apex of phallosoma.

Ecology
Specimens were collected in multistorey evergreen forests using fi sh traps.

Distribution
This species is only known from the type locality in Central Sulawesi and another locality in northern Sulawesi.

Remarks
A female specimen is chosen as the holotype for this species because the long, erect setae covering the body are not as obvious in the two male specimens available. One of the males was collected from the same collecting event as the females and shares all other diagnostic characters. The unique setation may be sexually dimorphic. Additional male specimens will be required to determine this. The male was dissected and the genitalia described here show that this species is very different from any other species of Physoderes.

Diagnosis
This species is recognized among congeners by the short and conical head, the scape not reaching the apex of the clypeus, the eye distinctly projecting, the submedian pronotal carina weakly defi ned, the apex of the scutellar process straw-colored, the parascutellar lobe rounded and skewed towards median, the corium dark brown with the membranous portion basally dark brown, including the external cell, and areas directly adjacent to the medial vein apical extension and fi rst anal vein apex with remaining membrane off-white and translucent, the abdominal dorsal surface dark brown, the apodeme of the anterior opening of the pygophore obsolete with the margin rounded, the cup-like sclerite with rounded apex and paired adjacent subacute protuberances and the dorsal phallothecal sclerite triangular in shape. This species is most similar to P. nigroalbus but can be differentiated from it by size, the coloration of the head, pronotum and scutellar process, and the shape of the eye.

Etymology
The name of this species is based on the Latin adjective "tricolor", meaning three-colored, to describe the three colors that are visible dorsally.
COLORATION (Figs 15, 18). Brown, brownish black and off-white. Head anteriorly straw-colored, posteriorly brown. Antenna straw-colored. First and second segment of labium straw-colored, third segment light brown. Anterior lobe of pronotum brown with straw-colored markings, posterior lobe brown, scutellum basally brown, apically straw-colored including scutellar process, pleuron straw-colored with brown patterns, sternum brown to dark brown. Corium of hemelytron dark brown, membrane basally dark brown, including the external cell, and areas directly adjacent to the medial vein apical extension and fi rst anal vein apex with remaining membrane off-white and translucent. Femora and tibiae of legs straw-colored with medial and apical brown annulations, tarsi and claws light brown. Abdomen dorsally dark brown, ventrally straw-colored with brown and dark brown patterns, anterior half of connexivum dark brown, posterior half straw-colored, exposed part of pygophore dark brown. VESTITURE. Sparsely setose. Head with some curved setae and short, fi ne, 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 irregular row of tuberculated, short, curved setae on lateral margins and curved setae on the anterior portion of dorsal surface, 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 not reaching apex of clypeus; eye distinctly projecting in dorsal view, less than 1/ 5 length of head.
THORAX. Antero-lateral paired projections acute, diverging; surface of anterior lobe smooth, ridges almost obsolete; median pronotal depression not contiguous with transverse sulcus; paramedian carina weakly defi ned; anterior pronotal lobe shorter and slightly narrower than posterior lobe, anterior lobe level with posterior lobe in lateral view; parascutellar lobe rounded, skewed towards median; scutellum rounded triangular, scutellar process long, apex subacute.
LEGS. Same as genus description.
GENITALIA. Anterior margin of mediosternite 8 undulating, without medial apodeme; transverse bridge of pygophore narrow, margin of anterior opening rounded, apodeme absent, apical margin of posterior opening smooth; cup-like sclerite apically rounded, with adjacent paired small subacute protuberances; arms of basal plate parallel to each other; ductifer membranous; endosomal struts tapered, with acute apex, divided into two arms basally; shape of dorsal phallothecal sclerite triangular with angular lateral prolongations.

Female
Similar to males except anterior pronotal lobe (2.17 mm average) distinctly narrower than posterior lobe (2.77 mm average) and median pronotal depression contiguous with transverse sulcus.

Ecology
Nothing is known about the biology of this species.

Distribution
This species is only known from the type locality of Sandakan, northeast Borneo.

Remarks
This species shares the general head and pronotal morphology of P. notata and is placed within the same clade as P. notata in the phylogenetic analysis above.

Physoderinae phylogeny: implications for biogeography and character evolution
Only three representatives of Physoderinae, two species of Physoderes and one of Breviphysoderes gen. nov., were previously included in a molecular phylogenetic analysis that also investigated divergence dates (Hwang & Weirauch 2012). In that analysis, Physoderinae diverged around 45 Mya from the Neotropical Nalata clade, their closest relatives included in that analysis. Divergences within the genus Physoderes were hypothesized to be as recent as 5 Mya. Given these timelines, dispersal, not vicariance, is likely the best explanation for the occurrence of Physoderinae in Madagascar. The nonmonophyletic nature of the Madagascan Physoderinae indicates that the evolutionary history of this group in the Madagascan region cannot be explained by a single colonization event with subsequent radiation. However, the majority of Madagascan Physoderinae are part of a single evolutionary lineage, Clade A, indicating that the morphological diversity found amongst Madagascan Physoderinae may in fact be due to a radiation in response to different ecological factors on the island. The single Australasian species of Paraphysoderes may then represent an "out of Madagascar" dispersal event, while the single Mauritian species of Physoderes could represent a second colonization from the Madagascan region. A formal biogeographic analysis based on morphological and molecular data that also includes divergence dating analyses is now the essential next step to test this proposed scenario.
Our analysis partially supports the hypothesis that the Neotropical species of Physoderinae that appear to have retained a number of plesiomorphic features as summarized by Weirauch (2006), are the sister group of all remaining Physoderinae: Leptophysoderes and Cryptophysoderes do in fact form a clade and are sister to all Australasian, Oriental, and Madagascan Physoderinae, but the Afrotropical Porcelloderes is recovered as the sister group to all remaining Physoderinae. This somewhat puzzling result may in part be derived from the fact that pronotal structures are strongly represented in our morphological dataset and the pronotum is highly modifi ed in the apterous Porcelloderes. This result is clearly in need of additional testing, preferably with the addition of molecular data, as is the apparent paraphyly of Leptophysoderes, which is likely due to the coding of two male (UCR_ENT 00014362, UCR_ENT 00014363) and two female (UCR_ENT 00014360, UCR_ENT 00014361) specimens that show signifi cant sexual dimorphism.
Despite being riddled by homoplasy, an examination of morphological characters allows for several observations. As an example, the highly modifi ed physoderines Tribelocephaloides, Henicocephaloides Villiers, 1962 and Mimoelasmodema, which are characterized by derived features including the ventral position of the eye and the lack of armature on the fore femur, are phylogenetically derived within their respective clades. In addition, unarmed and slender forelegs evolved independently from stout and heavily armed forelegs among the Neotropical clade (Leptophysoderes + Cryptophysoderes) and within a clade nested within the Madagascan Clade B (Befotaka Villiers, 1962+ Rodepirea Villiers, 1962+ Maroantsetrana Villiers, 1962. We speculate that this might be in response to a shift in prey organism or a different predatory strategy.

Comments on taxonomy
Based on fi ndings from our phylogenetic analysis that Physoderes is polyphyletic, we here redefi ne Physoderes and erect three new genera to accommodate the remaining species. We retain the name Physoderes for the clade that contains the type species of Physoderes, P. notata (Westwood, 1845), following the International Commission on Zoological Nomenclature. Redescriptions of Physoderes and Paraphysoderes and descriptions of the new genera are presented above. A summary of the taxonomic changes made in this publication and a list of new taxa described is also presented. A few species of Physoderes are placed as incertae sedis due to unknown locations of type specimens, leading to uncertainty in their identities. Attempts to locate these type specimens were unsuccessful, as the taxon authors did not specify depository sites and enquiries made to major European museums likely to house these specimens turned out negative.
Our revision of the Oriental and Australasian Physoderinae provides, for the fi rst time, a comprehensive picture of the diversity patterns in these regions. Notably, the highest number of species is found on two islands, Borneo and New Guinea. Borneo had already been identifi ed as a hotspot for physoderine species-level diversity by earlier workers (e.g., Miller 1940), but the addition of new species in the present study has emphasized this observation. Similarly, the addition of seven new species from New Guinea and the surrounding islands highlights the extreme diversity in this region. In contrast, the paucity of species described from the continental Oriental region may be an effect of poor sampling, a hypothesis that is supported by the fact that we discovered two new species from the Indian subcontinent for which we only examined a total of 42 specimens. Additional fi eld work in the continental Oriental region will likely reveal additional undescribed diversity.
Distributional data on physoderine species in the Oriental and Australasian regions do not provide a cohesive pattern with regards to factors that determine species ranges. Some species are widely distributed across island archipelagos, with insignifi cant morphological variation (see Physoderes curculionis, P. fuliginosa, P. notata). Other species appear to be highly restricted to a small endemic range close to their type locality, despite adequate sampling in surrounding regions and the fact that these species are not isolated on an oceanic island (see Physoderes muluensis sp. nov., P. minime sp. nov.). Further investigations into the natural history of the various physoderine species will be needed to explain the diversity patterns observed in the Oriental and Australasian regions. Measurements for species of Breviphysoderes gen. nov., Macrophysoderes gen. nov., Nanophysoderes gen. nov., Paraphysoderes Villiers, 1962 and Physoderes Westwood, 1945 (in mm