taxonID	type	description	language	source
03A38781FFE1FF87FF70FF7CFCA7C6C1.taxon	description	C. (C.) debaonica Xu, Tang & Skelley * C. (C.) collina Skelley, Xu & Tang, n. sp. C. (C.) lata (Grouvelle) C. (C.) fupingensis species group C. (C.) fupingensis Skelley, Tang & Xu C. (C.) nigra species group C. (C.) abyssa Skelley, Xu & Tang, n. sp. C. (C.) convexa Skelley, Xu & Tang, n. sp. C. (C.) cyclochasma Skelley, Xu & Tang, n. sp. C. (C.) discimaculata (Mader) C. (C.) eurynota Skelley, Xu & Tang, n. sp. C. (C.) intermedia (Chûjô) C. (C.) nigra (Gorham) C. (C.) torquata Skelley, Xu & Tang, n. sp. C. (C.) vittata (Arrow) C. (C.) yunnanensis (Grouvelle) C. (C.) papua species group C. (C.) papua Skelley, Xu & Tang, n. sp. C. (C.) samara Skelley, Xu & Tang, n. sp. Cycadophila (Strobilophila) Skelley, Xu & Tang, n. sg. C. (S.) assamensis Skelley, Xu & Tang, n. sp. C. (S.) hiepi Skelley, Xu & Tang, n. sp. C. (S.) kwaiensis Skelley, Xu & Tang, n. sp. C. (S.) tansachai Skelley, Xu & Tang, n. sp. * C. (S.) yangi Skelley, Xu & Tang, n. sp. The PCR product of the 16 S rRNA gene had a length of 311 – 313 bp. The alignment matrix was 317 bp long, with 5 gaps, 218 conserved sites, 96 variable sites, 67 parsimony-informative sites and 29 singleton sites. The NJ, MP and ML trees derived from DNA analyses were largely congruent with essentially the same terminal clades and no rearrangement of clades relative to one another. Only the one derived from ML is presented here (Fig. 1). This tree is similar to those presented by Xu et al. (2015), but is based on a larger sample of populations and species of Cycadophila and includes an additional (undescribed) taxon within the subfamily Pharaxonothinae as an outgroup. The same clades and branching patterns are present in our tree as in Xu et al. (2015), except that no members of the Strobilophila clade and papua clade were analyzed in the previous study. In the tree presented here Cycadophila + Pharaxonotha are a monophyletic grouping considered together in relation to the other members of the Pharaxonothinae sampled (bootstrap support = 100). Bootstrap support for the Pharaxonotha (sensu stricto) clade is also strong (> 90). Within this New World clade P. kirschii is sister to P. confusa, P. portophylla and P. floridana. With the addition of more populations and taxa to the analysis the genus Cycadophila remains monophyletic, although bootstrap support is relatively weak (<40). Within the subgenus Cycadophila this analysis recovered four strongly supported (bootstrap> 90) clades: 1) the lata species group (only C. debaonica and C. collina sampled); 2) the nigra species group (only C. yunnanensis and C. nigra sampled); 3) the fupingensis species group (consisting of a single species C. fupingensis); and 4) the papua species group (only C. samara sampled). The relationships among these four species groups remains unclear, since bootstrap support at nodes is relatively low (<40). For the subgenus Strobilophila, six of seven populations (three of four species) analyzed form a monophyletic group with one outlier [see more discussion below under Key to larvae and Remarks for C. (S). yangi]. Although bootstrap support for many nodes within the genus Cycadophila clade may be considered weak, our molecular data support our morphological hypotheses of relationships for this genus. Together molecular and morphological data support our concept of the genus Cycadophila as a monophyletic lineage.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEEFF87FF70FEA7FE8FC1B7.taxon	materials_examined	Type species. Cycadophila debaonica Xu, Tang & Skelley, 2015, by original designation.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEEFF87FF70FEA7FE8FC1B7.taxon	diagnosis	Adult diagnosis. The genus Cycadophila is easily distinguished from other genera of Pharaxonothinae by the head surface with transverse occipital ridge (vertexal line); large eyes that encroach upon head ventrally; supraocular striae present, often short; having a remnant of the submental-gular suture visible to variously depressed; male genitalia with median lobe and tegmen twisted, spiculum gastrale asymmetrical; and lacking abdominal calli. Many members of Cycadophila are known to be associated with Cycas spp. in SE Asia. Updated description. Length 3.04 – 6.95 mm, width 1.15 – 2.80 mm. Body form elongate, not strongly parallel sided; dorsoventrally flattened, weakly convex; widest anterior to or at middle of elytra; color pale to dark reddishbrown with or without variously sized dark macula on elytra; dorsal punctation fine, evenly distributed.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEEFF87FF70FEA7FE8FC1B7.taxon	description	Head with dorsal surface flat to convex (swollen) above eyes, clypeus flat or curved downward apically, may have vague depressions basally; clypeus apically truncate or rounded; supraocular striae present, extending along posterior 1 / 3 length of eye to entire length of eye; transverse occipital ridge present; stridulatory file on occipital region of head present, one on each side of the midline, widely separated, weakly convergent anteriorly; eyes prominent, coarsely faceted, large, ventrally encroaching upon gular area, distance between eye and lateral corner of mentum <1 / 2 eye length in ventral view; lacking lateral temple behind eye. Antennomeres I – VI length subequal to width, moniliform in most, slightly elongate in others; antenna with 3 - segmented club. Mouthparts with maxillary palp 4 - segmented, palpomere I very small, terminal maxillary palpomere cylindrical, acuminate apically, elongate, approximately 2 × longer than preceding palpomere; labial palp 3 - segmented; palpomere I small, elongate; II transverse; III large, flattened, roughly oval in shape, 3 / 4 length of terminal maxillary palpomere; mentum with large triangular base defined by carina; lateral pockets in front of carina not evident; submentum and gula flat, suture indicated centrally with pair of small pits often marked by dark coloration and a few punctures forming weak to strong U-shaped depression. Thorax. Pronotum with anterior marginal bead complete, fine; pronotal anterior margin near eyes weakly emarginate, anterior angles slightly projecting, angularly rounded; pronotal basal pits small, associated longitudinal groove fine, groove up to 1 / 4 length of pronotum; secondary transverse groove between pits present, curving along posterior margin; pronotal lateral margin evenly arcuate, lacking modifications; pronotal lateral marginal bead thin or thickened in both dorsal and lateral view. Prosternum in lateral view convex. Scutellum broadly pentagonal. Elytra finely setose, setae indistinct; with 10 complete striae of moderate puncture size; base of elytra with marginal bead. Prosternum mostly flat, depressed posterior of coxa; procoxal cavities narrowly open, procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; hypomeron with or without longitudinal striations. Protibia truncate and slightly to strongly dilated distally, without disto-lateral tooth; disto-ventral and lateral margins with row of distally directed fine spinules or ventrally directed stout spinules; meso- and metafemora and tibiae elongate, narrow to roundly dilated; mesofemur and tibia oval-dilated in some. Tarsi 5 - segmented, tarsomeres I – III pubescent ventrally with long, fine, dense, distally directed setae, not lobed beneath; tarsomere IV without dense ventral setae, short, narrow and inconspicuous. Sexual dimorphism present or absent, when present usually appearing as variously swollen or dilated tibiae or femora in males, which legs and segments involved are species specific. Abdomen with 5 visible ventrites, lacking coxal lines and internal calli. Male genitalia with median lobe and tegmen twisted like a corkscrew; median strut> 2 times longer than median lobe; flagellum shorter than median lobe, struts not coiled; spiculum gastrale asymmetrical. Note. Xu et al. (2015) provided additional images, some species accounts and key to genera of Pharaxonothinae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFECFF85FF70FD45FA4EC3DB.taxon	description	- Protibia weakly dilated distally with stout marginal spines on lateral apical 1 / 2 separated by their length or more (Figs. 2 G, 21 D); lacking or with greatly reduced furrow on each side of pronotal base (Fig. 21 A); Thailand .............................................................................................. Cycadophila (S.) yangi, new species 17 (16). Submental-gular suture indicated by weakly arcuate row of punctures; basal margin of pronotum nearly straight from side to side, weakly convex, lacking weak angular concavities where pronotal discal grooves join margin; scutellar shield more pentagonal; female with terminal abdominal ventrite convex apically, unmodified (Figs. 16 G, 19 G); male flagellum and penile struts longer, length of median lobe to penile strut 1: 5; Thailand, India .......................................... 18 - Submental-gular suture indicated by distinctly arcuate row of punctures; basal margin of pronotum sinuate, with distinct but shallow angular concavities where pronotal discal grooves join margin; scutellar shield more transverse; female with terminal abdominal ventrite concave apically, modified (Fig. 17 E); male flagellum and penile struts shorter, length of median lobe to penile strut 2: 7; Vietnam ............................................................ C. (S.) hiepi, new species 18 (17). Posterior lateral margin of pronotum straight to posterior angle which may have a very small indistinct projection behind pore (Fig. 18 A); no sexual dimorphism visible on terminal abdominal ventrite; inhabiting Cycas sp. on karst towers near the city of Kanchanaburi, Kanchanaburi province in Thailand at elevations below 100 m. ............ C. (S.) kwaiensis, new species - Posterior lateral margin of pronotum distinctly curved inward at posterior angle which has a distinct, small acute projection behind pore (Figs. 16 A, D, 19 A); females with terminal abdominal ventrite with a small area of fine punctures on apical medial surface, males evenly coarsely punctate throughout; occurring in widespread localities and hosts ..................... 19 19 (18). Elytral striae weakly impressed, giving intervals a weakly convex appearance, most evident on middle of disc; female terminal abdominal ventrite with apical medial surface glossy, glabrous and finely punctate, distinctly different from rest of surface, males with this surface entirely setose and coarsely punctate (Figs. 19 F – G) similar to rest of surface; inhabiting Cycas elephantipes, C. clivicola lutea, C. pectinata, C. petraea and C. tansachai above 300 m elevation in Thailand ............................................................................................ C. (S.) tansachai, new species - Elytral striae not impressed, intervals appearing flat; both male and female terminal abdominal ventrite with apical medial surface setose, females with finer punctures on apical medial surface than males (Figs. 16 F – G); inhabiting Cycas pectinata, Assam, India ................................................................ C. (S.) assamensis, new species	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEDFF81FF70FCF3FB32C75C.taxon	materials_examined	Type species. Cycadophila debaonica Xu, Tang & Skelley, 2015, by original designation.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEDFF81FF70FCF3FB32C75C.taxon	diagnosis	Adult diagnosis. Cycadophila (Cycadophila) is distinguished externally from the subgenus Strobilophila by the thin lateral pronotal carinae in lateral view with single row of punctation, by the lack of stout setae on the lateral margin of the protibia or with only a few setae at the extreme apex and with sexual dimorphism present or not, when present appearing on various legs. Adult description. Length 3.04 – 6.95 mm, width 1.15 – 2.80 mm. Body: form elongate, not strongly parallel sided; dorsoventrally flattened to weakly convex; widest at middle of elytra; color reddish-brown or dark brown, with or without color pattern; dorsal punctation moderately coarse, evenly distributed, dense. Head: dorsal surface of head relatively flat to convex in profile; convex between eyes, clypeus flat to convex apically; some with vague depressions basally; clypeus truncate anteriorly; supraocular striae length variable; transverse occipital ridge present (vertexal line); stridulatory file on occipital region of head present, one on each side, widely separated, parallel; eyes prominent, coarsely faceted; usually lacking lateral temple behind eye. Antenna: antennomere I – VI length subequal to width, moniliform in most; antenna with 3 - segmented club. Mouthparts: terminal maxillary palpomere cylindrical, acuminate at apex, elongate, 2 – 3 × longer than preceding palpomere; terminal labial palpomere flattened, roughly oval in shape, 3 / 4 length of terminal maxillary palpomere; mentum with large triangular base defined by carina; lateral pockets in front of carina evident or not; submentum and gula relatively flat, suture not obvious, medially with weak to distinct depression bearing setose punctures. Pronotum: pronotal anterior marginal bead complete, fine; pronotal anterior margin near eyes emarginate, anterior angles rounded; pronotal basal pits present, associated longitudinal groove distinct, groove approximately 1 / 4 pronotal length; secondary transverse groove between pits present, broad, curving along posterior margin except medially where slightly separated from margin; pronotal lateral carinae evenly arcuate, lacking modifications; pronotal lateral carinal bead narrow. Scutellar shield: broadly pentagonal. Elytra: finely setose, indistinct; with coarse, dense interval punctures; elytra with marginal bead basally. Prosternum: mostly flat, depressed or not posterior of coxa; procoxal cavities narrowly open; pronotal hypomeron with longitudinal striations, weak in some. Legs: protibia dilated distally or not, angled laterally but without apical tooth; ventral apical margin with or without fringe of stout ventrally directed, tooth-like spinules, some present on disto-lateral margin; meso- and metafemora elongate oval, meso- and metatibiae moderately angularly dilated at apex. Abdomen: ventrites I – IV each with pair of long erect sensory hairs located adjacent to midline (often abraded). Sexual dimorphism: when evident, present in variously developed legs of males, being more robust, dilated distally or variously curved. Male genitalia: median lobe and tegmen flattened laterally, weakly twisted basally; penile struts not fused, separate for entire length, each 5 – 6 × longer than median lobe; flagellum shorter than penile strut, not coiled; spiculum gastrale asymmetrical. Female genitalia: gonocoxites triangular, gradually narrowed posteriorly; apices of gonocoxites laterally with concave impression and setae. Gonostylus small, inserted at the concave impression of the gonocoxite, with several short setae and one long seta. Valvifers expanded posteriorly. Spermatheca elongated, C-shaped.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFEDFF81FF70FCF3FB32C75C.taxon	discussion	Remarks. Cycadophila (Cycadophila) presently comprises four species groups as diagnosed below: the lata group, fupingensis group, nigra group and papua group. Preliminary morphological and molecular data show these to be monophyletic groups, but further data and analyses are needed to substantiate relationships amongst them before they can be confidently assigned into separate subgenera. They are readily distinguished from each other by characters in the key and by diagnostic characters presented in the following accounts. The external morphological diversity of these groups makes a simple diagnosis of C. (Cycadophila) as presented here difficult.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF81FF70FE53FEFAC20A.taxon	diagnosis	Adult diagnosis. The Cycadophila (Cycadophila) lata species group is distinguished from other members of Cycadophila by color: body reddish-brown with dark mark along elytral suture; head dorsal surface flattened in profile; supraocular stria present, extending full length of eye; maxillary palpi normal in length, not more than 2 × longer than labial palpi; ventral interocular distance <1 / 3 head width; submental-gular suture primarily flattened, medially with weak, darkened transverse depression with row of setose punctures; pronotal anterior angles rounded, surface near anterior angle convex; lateral carinae evenly arched most of length, basally appearing evenly sinuate; pronotal lateral carina thin in lateral view, with single row of setose punctuation on marginal surface; prosternal process narrow (<1 / 2 width of profemur), rounded at apex, apex depressed, in lateral view convex; protibia dilated distally, with fringe of stout setae along apical margin and a few at extreme apical-lateral margins; coxal line behind mesocoxae without punctures; male genitalia with tegmen strongly twisted basally, penile strut 3 × length of median lobe; sexual dimorphism present with males having strongly dilated pro- and mesotibiae and femora.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF81FF70FE53FEFAC20A.taxon	discussion	Remarks. This species group contains three species that are similar in morphology. Current molecular data indicate the available specimens of this group to be part of a complex of species. Here we only present accounts of the two described species and one new species belonging to this group. Further work is underway to help resolve this complex.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF9EFF70FB6AFE55C564.taxon	diagnosis	Adult diagnosis. Morphologically distinguishable from other C. (Cycadophila) spp. by the long supraocular striae, head width / ventral interocular distance ratio <3.50; width of elytra greatest at middle, without fine acute projection at posterior pronotal hind angle, and strong distal dilation of mesotibia and mesofemora in major males. Morphologically most similar to C. (C.) debaonica which has a fine obtuse projection at posterior pronotal hind angle and major males with mesofemora narrowing from middle to apex. Most confidently distinguished from C. (C.) debaonica by analysis of the 16 S rRNA gene (GenBank Accession numbers KY 365227 and KY 365247), host and distribution. Adult male description. Length 4.19 – 5.85 mm, width 1.88 – 2.40 mm. Body elongate-oval in dorsal view, greatest width at middle of elytra; in lateral view convex dorsally. General body color reddish-brown (Figs. 4 A – D), head darker than pronotum and lateral halves of elytra; elytra with large central black maculation occupying roughly half of elytral length from near the scutellum to 2 / 3 length of elytra and roughly 2 / 5 of the elytral width, dorsal surface punctate, shining and dorsally appearing glabrous, short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF9EFF70FB6AFE55C564.taxon	description	Head in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, finely, sparsely punctured; width 1.12 mm; dorsal interocular distance 0.68 mm, head width / dorsal interocular distance ratio 1.65, ventral interocular distance 0.39 mm, head width / ventral interocular distance ratio 3.05. Eye globular; bordered dorsally with long supraocular stria reaching anterior angle of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin anterior to antennal insertion. Antennal length slightly shorter than pronotal width, slightly longer than head width, antennomere relative lengths from base to end approximately 14: 10: 12: 10: 10: 10: 9: 9: 10: 10: 10, antennomere widths of II – VII subequal, I and VIII subequal and about 1.30 × width of II – VII, width equals length; IX and X subequal and about 2.00 × width of II – VII, XI about 1.50 × width of II – VII, globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Mandible with 3 apical teeth; prostheca extending from mola halfway to apical teeth, length about ½ width, edge nearly straight, with fringe of hair along dorsal base that extends to the proximal apical tooth; mola with carinulation and small spines. Labrum transverse, about 2.00 × broader than long; anterior margin fringed with long bristles, especially dense medially. Hypopharynx with thin setae directed medially. Maxilla with cardo expanded apically; stipes narrowly triangular, apically acute, partially overlapping with palpiger; galea and lacinia with long, densely aligned setae; setae on the lacinia thick and strong, but distinguishable hooks absent from surrounding dense setae; galea wider than lacinia. Maxillary palpomeres II, III and IV longer than width; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 5: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum, where shallow transversely elongate foveae are present bearing dense setose punctures (Fig. 4 D), foveae dark brown. Thorax with pronotum transverse in dorsal view with marginal beads on all sides; length / width ratio 0.69, convex, wider at base; anterior angles weakly developed; posterior angles exteriorly angulate; posterior margin projecting medially; with a narrow longitudinal furrow posterolaterally, beginning at pore on posterior margin extending forward and ending abruptly, length of furrows relative to pronotal length 0.22. Prosternum with anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae; prosternal process round and convex apically. Hypomeron laterally with few punctures, medially without longitudinal striations. Scutellar shield transverse, posterior margin clearly projected, pentagonal. Elytra in dorsal view elongate-oval, convex; length / width 3.30, greatest width near midlength; with marginal line basally, scutellary striole extending ¼ elytral length, with 10 – 13 punctures; intervals of striae with fine, shallow punctures. All punctures of elytral striae bearing a single short seta; seta only visible in profile, extending slightly out of puncture. Punctation on meso- and metaventrites shallow, small. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout, relatively similar in length and shape. Femora compressed laterally; tibiae shorter than femora, gradually dilated to narrow apex; protibia distinctly dilated apically, with apical fringe of short, stout spinules ventrally, inner margin with fine setae. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair of long erect sensory hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short, appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 90 ˚ toward the left side of the animal and resting on side. Tegmen sclerotized, triangular, but twisted medially; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres apically with long setae; length / width ratio 2.60. Aedeagus with median lobe slender, highly sclerotized, apex long and acuminate. Ratio of penile strut length to median lobe length 3: 1. Female similar to male except with pro- and mesotibiae and femora less robust and not dilated apically, pygidium truncate, apical edge fairly straight with 7 long setae and a fringe of setae about 1 / 2 as long. Genitalia with gonocoxites triangular, gradually narrowing posteriorly; apices of gonocoxites laterally with concave impression and setae. Gonostylus inserted at the concave impression of the gonocoxite, with several short setae and 1 long seta. Valvifers expanded posteriorly. Spermatheca elongate, C-shaped. Variation. In some individuals the dark elytral mark is slightly larger or smaller, and varies from a lighter to darker brown. In some this mark is pale enough to be indistinct. Variation in pro- and mesotibial dilation appears to be sexually dimorphic. Males usually have these tibiae more strongly dilated. However, there is enough variation that distinguishing weakly developed males from strongly developed females can be problematic. Weakly developed females have the mesotibia almost parallel-sided to the apex. Strongly developed males have the outer margin of the mesotibia slightly swollen to a widened apex. Similarly, mesofemoral development varies in males from strongly to weakly dilated toward apex. Major males of C. collina and C. debaonica are distinctly different in this character. Minor males of either species are difficult to distinguish. Figures used in the key are major males. Type locality. Laos, Houphan Province, Viengxay District, Vieng Xai village, 20 ˚ 23 ’ 41 ” N 104 ˚ 13 ’ 44 ” E. Range. Known from Houphan Province in northern Laos and Son La Province, Vietnam on Cycas collina, and the southern portion of Yunnan Province, China on Cycas sp. aff. collina and C. tanqingii.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF9EFF70FB6AFE55C564.taxon	materials_examined	Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ LAOS, Houphan Pr., Viengxay Ds., Vieng Xai vill., 20 ° 23 ' 41 ″ N, 104 ° 13 ' 44 ″ E, 7 - IV- 2015, N. T. Hiep, L. Averyanov, N. S. Khang, N. Q. Hieu, T. Maisak, Somneux ”; 2) “ ex ♂ cones Cycas collina (LA-VN 1124) ”; 3) [rectangular; red, printed in black ink] “ HOLOTYPE ♂ Cycadophila collina P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA. Allotype female and 45 paratypes: CHINA, Yunnan Province, Guanping Reserve, 22 ° 10 ' N 100 ° 50 ' E, 10 May 1996, W. Tang, ex spent ♂ strobilus of Cycas sp. aff. collina, WT 44 (2); ex early pollen-shedding ♂ strobilus of Cycas sp. aff. collina, WT 45 (allotype, FSCA; 20); ex receptive ♀ strobilus of Cycas sp. aff. collina, WT 46 (2); VIETNAM, Son La Province, nr. Mai Son, 21 ° 8 ' 630 ″ N 104 ° 1 ' 731 ″ E, Cycas collina, Apr 4, 1996, S. L. Yang SLY 777 (21). Paratypes deposited in ANIC, FSCA. Others Examined: China, Yunnan Province, Luchun Co, Cycas tanqingii ♂ cone, 27 March 2008, He Jianghai [specimen destroyed during DNA analysis; pin and label remain].	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF9EFF70FB6AFE55C564.taxon	etymology	Etymology. The species is named after one of its host species, Cycas collina.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFE8FF9EFF70FB6AFE55C564.taxon	discussion	Remarks. DNA analysis of the 16 S rRNA gene indicates that Cycadophila collina forms part of a cryptic species complex with C. debonica and that this beetle is associated with two hosts, Cycas tanqingii and the Cycas collina species complex. These two host Cycas species inhabit rugged terrain south of the Red River while the two known hosts of its sister taxon, Cycadophila debaonica, occur either far north of the Red River (Cycas debaoensis) or farther east and south of the Red River on limestone towers (Cycas hoabinhensis). Other possible host Cycas species for Cycadophila collina include Cycas dolichophylla, C. bifida and C. fugax, which occur in regions just north or east of Cycas collina and C. tanqingii (Osborne et al. 2007). Morphologically, C. collina is most similar to C. lata in development of male sexual dimorphisms. These species also appear to occupy distinct, but neighboring distributions (see Remarks under C. lata). Even without molecular analyses, the striking difference in ventral interocular distance and different distributions allow us to maintain C. lata and C. collina as valid species within the species complex. Considering the morphological similarity in species of the lata complex, the Laos holotype of C. collina was chosen because it was distinguished with both molecular and morphological evidence. Although the series of Chinese and Vietnamese paratypes match the Laos specimen morphologically, they have not been confirmed with 16 S rRNA evidence.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF7FF9EFF70FC44FF28C1EF.taxon	diagnosis	Adult diagnosis. A member of the C. (Cycadophila) lata species group as discussed above, distinguished most readily by the long supraocular striae, head width / ventral interocular distance ratio <3.5; width of elytra greatest at middle, with fine actute projection at posterior pronotal hind angle, and major males with mesofemora gradually narrowing to the apex. Most similar morphologically to C. collina which has an indistinct obtuse projection at posterior pronotal hind angle and major males with mesofemora parallel-sided nearly to apex. Most confidently distinguishable from C. (C.) collina by analysis of the 16 S rRNA gene (GenBank Accession numbers KR 005715 – KR 00717, KY 365239, KY 365249, KY 365251), as discussed earlier. Full descriptions and images of adults and larvae are presented in Xu et al. (2015).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF7FF9EFF70FC44FF28C1EF.taxon	materials_examined	Type locality. China, Guangxi Province, Debao County, Fuping village, 23 ° 29.624 ' N, 106 ° 12.980 ' E. Range. Known from Debao and Napo counties of Guangxi Province, China. The range of the host, C. debaoensis, extends to Banshui, Baise City, Guangxi Province and Funing County, Yunnan Province with 16 known natural populations (Xie et al. 2005; Fang 2009). Material examined. Holotype, allotype and paratypes as discussed in Xu et al. (2015). Others examined: VIETNAM, Hoa Binh Province, Lac Thuy Distr., Phu Lao Comm., 20 ° 33 ' 19 ″ N, 105 ° 45 ' 42 ″ E, 50 – 100 m, 25 - IV- 2015, N. S. Khang ex ♂ Cycas hoabinhensis (1; deposited at the FSCA).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF7FF9EFF70FC44FF28C1EF.taxon	discussion	Remarks. Xu et al. (2015) demonstrated that adults and larvae of this species occur together in male cones of Cycas debaoensis in Guangxi, China, where they appear to feed and reproduce. DNA analysis of the 16 S rRNA gene indicates that a population of Cycadophila debaonica in Vietnam south of Hanoi on Cycas hoabinhensis display DNA sequences similar to those type populations in Guangxi Province, China. Populations in northern Vietnam west of Hanoi, in northern Laos and in southern Yunnan Province, China on Cycas tanqingii and the C. collina species complex exhibit enough genetic differences to warrant recognition as a species distinct from Cycadophila debaonica and is named above as Cycadophila collina. Morphologically, many specimens of these two species are difficult to distinguish and form part of a cryptic species complex. They are both, however, morphologically distinguishable from the closely related Cycadophila lata (discussed below), which also occurs in Vietnam.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF4FF9BFF70FF7CFD4DC59E.taxon	diagnosis	Adult diagnosis. A member of the C. (Cycadophila) lata species group, with long supraocular striae, distinguished most readily from C. debaonica and C. collina by the head width / ventral interocular distance ratio> 4.0 0; width of elytra greatest anterior to middle, and their tibial dilation. Mesotibial and femoral dilation in the male holotype of C. lata is strong (Fig. 2 J), indicating it is a major male. Only the male holotype is known, females and minor males are unknown. Adult description. Length 4.98 mm, width 1.94 mm. Body in dorsal view elongate-oval, greatest width anterior to middle of elytra; in lateral view convex dorsally. General body color reddish-brown, head darker, slightly darker than pronotum and lateral halves of elytra, elytra with large central black maculation occupying half of elytral length; dorsal surface punctate, shining and dorsally appearing glabrous, short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF4FF9BFF70FF7CFD4DC59E.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex; finely, sparsely punctured; width 1.32 mm; dorsal interocular distance 0.56 mm, head width / dorsal interocular distance ratio 2.35, ventral interocular distance 0.29 mm, head width / ventral interocular distance ratio 4.50. Eye globular; bordered dorsally with long supraocular stria reaching anterior angle of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length slightly shorter than pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 16: 10: 11: 10: 10: 10: 10: 9: 11: 11: 16; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX and X similar in length, XI slightly longer, globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Mouthparts not dissected on unique holotype. Maxillary palpomeres II, III and IV longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palp III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with narrow marginal beads on all sides; length / width ratio 0.63, convex, wider basally; anterior angles weakly developed; posterior angle exteriorly angulate; posterior margin projecting medially; with narrow longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.25. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron without longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytral color pattern with dark macula on each elytron extending from suture to 2 / 5 width of the elytron and from near the scutellum to 2 / 3 length of elytra; in dorsal view elongate-oval, convex; greatest width near midlength; with marginal bead basally, scutellary striole with 11 punctures along anterior 1 / 4; intervals of striae with fine and shallow punctures. All elytral punctures bearing a single short seta; seta only visible in profile, extending slightly out of puncture. Punctation on meso- and metaventrite shallow and small. Metaventrite convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout; femora compressed laterally; tibiae shorter than femora, gradually dilated to narrow apex, with lateral margins convexly curved, most notable on mesotibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair of long hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 90 ˚ toward the left side of animal and resting on side. Tegmen sclerotized, triangular, but twisted in the middle; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.00. Aedeagus with median lobe slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 2. Female unrecognized.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF4FF9BFF70FF7CFD4DC59E.taxon	materials_examined	Type locality. Tonkin, Than moi. Range. Tonkin, Than moi. Material examined. Holotype male with the following labels (Fig. 5 E): 1) [rectangular; white; hand written in black ink] “ Tonkin, Than moi, 3 - 06, Coll. Fouquet ”; 2) [rectangular; red, printed in black ink] “ Type ”; 3) [rectangular; white; hand written in black ink] “ Pharaxonotha lata Grouv. ”; 4) [rectangular; white, printed in black ink] “ Erotylidae: Cycadophila lata (Grouv.) det. P. Skelley 2012 ”. Deposited in MNHN (examined).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF4FF9BFF70FF7CFD4DC59E.taxon	discussion	Remarks. Type locality of “ Tonkin ” corresponds to northern Vietnam and adjacent parts of Laos. There are approximately 17 species of Cycas (Osborne et al. 2007; Averyanov et al. 2014) occurring naturally in this area occupying a variety of habitats including cliff faces, forest understory and seashore. The collection locality for the holotype, interpreted here as “ Than moi ” (see Fig. 5 E) corresponds most closely to the present day locality of Thanh Moi (21 ° 38 ' N, 106 ° 33 ' E) in Long Son Province, Vietnam along Highway 1 A, built by the French in the early 20 th century. This is the habitat zone for Cycas ferruginea, a cliff-dwelling species (Osborne et al. 2007; Tang, unpub. obs.). Other less probable hosts occurring in the surrounding region are the understory forest cycads C. balansae, C. bifida and C. dolichophylla. Another possible candidate place name is Thanh Mai corresponding to two localities in Vietnam (20 ° 38 ' N, 106 ° 27 ' E or 21 ° 12 ' N, 106 ° 27 ' E) not located near any major roads, and therefore less likely to have been the original collecting site. These are situated further to the southeast in two adjoining provinces, Thai Binh and Hai Dong. The most likely host in this area is Cycas haobinhensis (Osborne et al. 2007), a species restricted to limestone cliff faces, which is already known to harbor a congener, Cycadophila debaonica (see above under Remarks for C. debaonica). Based on variation in other species of this group, the holotype of C. lata appears to be a major male and is the only known specimen. This prevents us from observing any variation, confirming female characters, or comparing molecular data. Further analysis must wait until potentially conspecific specimens are available for study.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF2FF9BFF70FC10FBC5C0A3.taxon	diagnosis	Adult diagnosis. The C. (Cycadophila) fupingensis species group is distinguished from other members of Cycadophila by the reddish-brown body with dark mark along elytral suture; head dorsal surface flattened in profile; supraocular stria present, short, extending basal 1 / 3 length of eye; maxillary palpi normal in length, not more than 2 × longer than labial palpi; ventral interocular distance approximately 1 / 3 head width; submental-gular suture primarily flattened, medially with weak, darkened transverse depression with row of setose punctures; pronotal anterior angles rounded, surface near anterior angle convex; lateral carinae evenly arched most of length, basal margin appearing evenly sinuate; pronotal lateral carina thin in lateral view, with single row of setose punctation on marginal surface; prosternal process narrow (<1 / 2 width of profemur), truncated at apex, apex depressed, in lateral view convex; protibia weakly dilated distally, lacking fringe of stout setae along apical and lateral margins; coxal line behind mesocoxae without punctures; male genitalia with tegmen moderately twisted basally, penile strut 5.0 0 × length of median lobe; and sexual dimorphism present with males having strongly dilated pro- and mesotibiae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF2FF9BFF70FC10FBC5C0A3.taxon	discussion	Remarks. Cycadophila fupingensis, currently the only member in the group, has morphological similarities with other groups in the subgenus. For example, the flattened head and general body shape are most similar to the lata group, the short ocular stria is most similar to the nigra group, and the narrowed tibia lacking stout setae with no notable dimorphism in on the legs and the male genitalia are most similar to the papua group. Molecular data analyzed in this paper place this group closest to the lata species group (Fig. 1).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF2FF9AFF70F906FD24C491.taxon	diagnosis	Adult diagnosis. A member of the C. (Cycadophila) fupingensis species group as discussed above, distinguished from other species of the genus by the narrow protibia lacking stout setae apically, short supraocular stria, fine pronotal punctation, and having a small elytral maculation. Full descriptions and images of adults and larvae are presented in Xu et al. (2015).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF2FF9AFF70F906FD24C491.taxon	materials_examined	Type locality. China, Guangxi Province, Debao County, Fuping village, 23 ° 29.624 ' N, 106 ° 12.980 ' E. Range. Known from Debao and Napo Counties of Guangxi Province, China on Cycas debaoensis. The range of this host, C. debaoensis, extends to Banshui, Baise City, Guangxi Province and Funing County, Yunnan Province with 16 known natural populations (Xie et al. 2005; Fang 2009). In Vietnam it occurs in Hoa Binh Province, Lac Thuy Distr. on C. hoabinhensis. Material examined. Holotype, allotype and paratypes as stated in Xu et al. (2015). Others examined: CHINA, Yunnan Province, Red River Valley, nr. Man Hao, 23 ° 01 ' N, 103 ° 24 ' E, 955 m, 12 May 1996, W. Tang, ex near receptive ♀ strobilus of Cycas diannanensis (type locality), WT 50 (2); Guilinchin Town, near Jingping, 22 ° 46 ' N, 103 ° 15 ' E, 9 May 1996, S. - L. Yang, ex ♂ strobilus of Cycas dolichopylla, WT 52 (4). VIETNAM, Hoa Binh Province, Lac Thuy Distr., Phu Lao Comm., 20 ° 33 ' 19 ″ N, 105 ° 45 ' 42 ″ E, 50 – 100 m, 25 - IV- 2015, N. S. Khang ex ♂ Cycas hoabinhensis (1). These are deposited at ANIC and FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF2FF9AFF70F906FD24C491.taxon	discussion	Remarks. Xu et al. (2015) demonstrated that adults and larvae occur together in male cones of Cycas debaoensis, where they appear to feed and reproduce.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF3FF9AFF70FD17FDD1C1AA.taxon	diagnosis	Adult diagnosis. The Cycadophila (Cycadophila) nigra species group is easily distinguished from other members of Cycadophila by the dark brown to black body often with elytral markings; head dorsal surface convex in profile, most notable over frons and clypeus; supraocular stria present, short, <1 / 2 posterior length of eye; maxillary palpi normal in length, not more than 2 × longer than labial palpi; ventral interocular distance approximately 1 / 2 head width; submental-gular suture primarily flattened, medially with depression between two pits, depression and pits variable in depth from just notable to sharply defined; pronotal angles sharply angulate, surface near anterior angle usually more convex (reflexed), lateral carinae straight most of middle length, basal margin appearing quadrate, straight at sides, angulate next to juncture with discal groove and pore, then evenly rounded (lobed) at middle; pronotal lateral carina thin in lateral view, with single row of setose punctation on marginal surface; prosternal process broad (> 1 / 2 width of profemur), truncated at apex, apex not notably depressed, in lateral view flattened; protibia weakly dilated distally, lacking fringe of stout setae along apical and lateral margins, may have couple of stout setae at disto-lateral angle; coxal line behind mesocoxae usually with large deep punctures; male genitalia with tegmen moderately twisted basally, penile strut 5.00 – 6. 0 0 × length of median lobe; and sexual dimorphism when present involves some modification to the male hind leg.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF3FF9AFF70FD17FDD1C1AA.taxon	discussion	Remarks. Some members of the nigra species group are superficially similar in general appearance to Pharaxonotha kirschii (a New World species, occasionally reported as a stored product pest, the Mexican grain beetle; Hinton 1945; USDA 2015). The similarity is so striking that entirely dark specimens of the nigra group can be easily confused with P. kirschii. However, the presence of a small temple behind the eye, dorsoventrally flattened male genitalia, male with small tooth on apex of last abdominal ventrite, and distribution readily distinguish P. kirschii from the nigra group which mostly lack the temple, have twisted genitalia, and lack sexually dimorphic abdominal ventrites. The striking similarity in general body plans between the cycad-inhabiting members of Pharaxonotha and those of Cycadophila other than the nigra species group, may be convergences based on similarities in life histories on cycads. In contrast, the superficial similarity of P. kirschii (the only Pharaxonotha sp. not associated with cycads) and those of the nigra species group, may indicate similarities in life histories not involving cycad cones. The general paucity of some of these species in available materials collected on cycads supports the hypothesis that some members of the nigra species group are not obligate cycad cone associates and may feed on other hosts. The nigra species group is the most diverse group of Cycadophila. Molecular analyses of two species are presented in this paper and they are consistent with the concept of a monophyletic clade, but the variability in morphological characters of taxa lacking molecular data do not yet warrant naming of this group as a distinct subgenus or to split it up further.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF0FF97FF70FF7CFAFAC795.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above readily distinguished from all others by its large body size, large deep submental-gular pit, and transverse pronotum. Adult description. Length 4.67 – 6.95 mm, width 1.73 – 2.50 mm. Body in dorsal view elongate, sides gradually arched, almost parallel-sided, greatest width at middle of elytra; in lateral view convex dorsally. General body color entirely dark reddish-brown, appendages and lateral margins lighter; dorsal surface distinctly punctate, surface appearing slightly shiny, dorsal punctation with short procumbent hairs barely protruding out of punctures, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF0FF97FF70FF7CFAFAC795.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface convex, appearing swollen above eye, distinctly coarsely punctured; width 1.00 – 1.27 mm; dorsal interocular distance 0.62 – 0.80 mm, head width / dorsal interocular distance ratio 1.60 – 1.62, ventral interocular distance 0.52 – 0.64 mm, head width / ventral interocular distance ratio 1.91 – 2.00. Eye globular; bordered dorsally by swollen head, so that dorsal margin of eye with deep groove, supraocular stria less than half length as eye but coarsely punctured; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 17: 11: 12: 10: 10: 10: 10: 10: 14: 14: 17; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly shorter than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus rounded apically, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate, apex densely papillate; relative lengths of II – IV approximately 4: 2: 6. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Submentum with coarse punctures distinct, separated by glossy cuticle. Gular area smooth, without setae or punctation, except at suture with submentum where a deep circular depression surrounds a central area with setose punctures; gula with deep groove surrounding medial and posterior eye margins. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.73 – 0.79, convex, nearly parallel-sided, vaguely narrowing anteriorly; anterior angles projecting, weakly angulate; lateral carinae gradually turning inward anteriorly and posteriorly, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 °; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.20 – 0.22. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate, not turning inward at apex. Hypomeron with coarse punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin roundly projected, pentagonal. Elytra elongate with sides gradually arched, convex; length / width 3.41 – 3.65, greatest width near midlength; with marginal bead basally, scutellary striole with 9 – 10 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta length just reaching outside of punctures. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length; line behind mesocoxae with distinct, coarse punctures. Legs narrow, similar in length; femora compressed laterally; male with metafemur swollen basally, posterior margin curved; tibiae shorter than femora, weakly dilated to narrow apex. Protibia not dilated at apex, with fine setal fringe on ventral, lateral margin smooth without stout spinules; few small stout spinules at disto-lateral angle. Meso- and metatibia narrow, similar to protibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV lacking pair of erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side (Fig. 6 E). Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat spatula-shaped parameres. Parameres with long setae apically; length / width ratio 2.00. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 2: 9. Female similar to male except for narrow metafemur with evenly curved posterior edge	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF0FF97FF70FF7CFAFAC795.taxon	materials_examined	Type locality. CHINA: Hong Kong Isl., Jardine’s Lookout. Range. China (Hong Kong, Hainan Prov.). Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ CHINA: Hong Kong Isl., Jardine’s Lookout, 6 April 2000, fruit trap, Leg. J. Mat; é ” 2) [rectangular; red, printed in black ink] “ HOLOTYPE ♂ Cycadophila abyssa P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA. Allotype female and 7 paratypes: CHINA: Hong Kong Isl., Jardine’s Lookout, 6 April 2000, fruit trap, Leg. J. Maté (allotype, FSCA, 4); Hong Kong [on reverse] 48 29; [second label] 547 (1); CHINA: Hainan, ex cone Cycas hainanensis, V- 2001, Ma Xaio Yan, 3 - 1 - 2, Photo Code XB- 1 (1); CHINA: Hainan Prov., Wuzhishan City, Wushishan Mt.; 700 – 1000 m, 15 ° 54 ' N, 109 ° 41 ' E, 18 - IV- 2012, YIN Ziwei (1). Paratypes deposited in BMNH, FSCA, NZAC, SNUC.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF0FF97FF70FF7CFAFAC795.taxon	etymology	Etymology. The specific epithet is derived from the Greek ‘ abyssos’ meaning bottomless pit, used in reference to the pit at the submental-gular juncture.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF0FF97FF70FF7CFAFAC795.taxon	discussion	Remarks. One paratype was collected on a cycad cone, indicating a possible cycad association. Wild Cycas populations in the section Stangerioides subsection Taiwanianae occur in the vicinity of all type localities of this species, including C. fairylakea in Shenzhen at the border with Hong Kong and C. hainaensis on Hainan Island. Additional materials are needed to substantiate this relationship. See remarks under C. (C.) cyclochasma.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFEFF95FF70FE14FC57C246.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above distinguished by the narrow protibia lacking stout setae apically, short supraocular stria, pronotum with anterior margin straight and disc evenly convex to lateral margins, elytra with distinct color pattern, and male with enlarged metafemur and curved metatibia. Adult description. Length 3.04 – 3.46 mm, width 1.15 – 1.31 mm. Body in dorsal view elongate, sides arcuate, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color dark brown, appendages paler; elytra primarily yellow-brown with dark maculation along suture from basal to 2 / 3 length, notably wider along middle, and with dark mark near base of lateral margin; dorsal surface distinctly punctate, surface shiny, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes, elytral setae very short, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFEFF95FF70FE14FC57C246.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface convex, distinctly punctured; width 0.60 – 0.67 mm; dorsal interocular distance 0.40 – 0.44 mm, head width / dorsal interocular distance ratio 1.52 – 1.53, ventral interocular distance 0.28 – 0.30 mm, head width / ventral interocular distance ratio 2.17 – 2.23. Eye globular; bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 18: 14: 15: 10: 10: 10: 10: 10: 15: 15: 22; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a small depression is present with setose punctures. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.75 – 0.76, surface convex to lateral carinae; anterior angles rounded; lateral carina arched anteriorly, straight basally, with thin marginal bead, bead bearing row of minutely setose punctures; posterior angle exteriorly angulate; posterior margin straight laterally, weakly lobed medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.20 – 0.21. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate and flat in lateral view, weakly projecting. Hypomeron with few punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin rounded, somewhat pentagonal. Elytra elongate, evenly arcuate along sides, convex; length / width 3.33 – 3.35, greatest width near midlength; with marginal bead basally, scutellary striole with 10 – 12 punctures; intervals of striae with fine, distinct punctures. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, metathoracic discrimen indistinct, extending slightly over half metaventrite length. Legs narrow, similar in length; femora compressed laterally, enlarged with curved sides; tibiae shorter than femora, narrow at apex. Protibia narrow, weakly angled on lateral apex, apex fringed with small, hair-like spinules, lateral margin lacking spinules, couple stout spinules present on apical-lateral angle. Meso- and metatibia narrowed, obliquely angled at lateral apex. Male metafemur enlarged, club-like with distinctly curved posterior margin; metatibia curved with slight rounded swelling on inner apical half. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV apparently lacking erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres short, with long setae apically; length / width ratio 2.00. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 4. Female similar to male except legs narrower, metafemur not as enlarged, metatibia not curved.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFEFF95FF70FE14FC57C246.taxon	materials_examined	Type locality. Thailand, Kaen Municipality. Range. Thailand, Kaen Municipality. The word for municipality in Thai is “ Muang ” and “ Muang Kaen ” is the name widely used in Thailand for Kanchanaburi province, so Kanchanaburi province is the probable origin. During this study, populations of three species of Cycas were surveyed in Kanchanaburi province, including C. siamensis, C. pectinata and a third undescribed species. None yielded specimens matching Cycadophila convexa. Another possible locality is Khon Kaen, which is known to contain one Cycas species, C. siamensis. Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ THAILAND: Kaen Municipality, 9 - V- 1954, R. E. Elbel ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila convexa P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the USNM. Allotype female and 3 male paratypes: THAILAND: Kaen Municipality, 9 - V- 1954, R. E. Elbel (holotype & allotype, USNM, 3); 8 - V- 1954 (1); 12 - V- 1954 (1). Paratypes deposited in USNM, FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFEFF95FF70FE14FC57C246.taxon	etymology	Etymology. The species is named for the unique pronotal disc which is completely convex all the way to the lateral margins.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFEFF95FF70FE14FC57C246.taxon	discussion	Remarks. The straight anterior pronotal margin of C. convexa is similar only with that of C. torquata. Despite surveys in likely localities and hosts (see comments under “ Range ” above), this species has not been collected with Cycas, suggesting that it does not have an association with cycads.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFCFF93FF70FB26FBD1C06A.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above readily distinguished from all others by its large body size, large deep submental-gular pit, and elongate pronotum with nearly straight sides. Adult description. Length 6.90 mm, width 2.50 mm. Body in dorsal view elongate, sides gradually arched, almost parallel-sided, greatest width at middle of elytra; in lateral view convex dorsally. General body color entirely dark reddish-brown, appendages and lateral margins lighter; dorsal surface distinctly punctate, surface appearing slightly shiny, dorsal punctation with short procumbent hairs barely protruding out of punctures, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFCFF93FF70FB26FBD1C06A.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface convex, appearing swollen above eye, distinctly coarsely punctured; width 1.05 mm; dorsal interocular distance 0.67 mm, head width / dorsal interocular distance ratio 1.57, ventral interocular distance 0.55 mm, head width / ventral interocular distance ratio 1.91. Eye globular; bordered dorsally by swollen head, so that dorsal margin of eye with deep groove, supraocular stria less than half length of eye but coarsely punctured; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 16: 11: 13: 10: 10: 10: 10: 11: 16: 16: 17; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly shorter than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus rounded apically, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate, apex densely papillate; relative lengths of II – IV approximately 4: 2: 6. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except at suture with submentum where a deep circular depression surrounds a central area with setose punctures; gula with deep groove surrounding medial and posterior eye margins. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.82, convex, nearly parallel-sided, vaguely narrowing anteriorly; anterior angles projecting, weakly angulate; lateral carinae nearly straight, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 o; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.25. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate, not turning inward at apex. Hypomeron with coarse punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin roundly projected, pentagonal. Elytra elongate with sides gradually arched, convex; length / width 3.52, greatest width near midlength; with marginal bead basally, scutellary striole with 9 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta length just reaching outside of punctures. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length; line behind mesocoxae with distinct, coarse punctures. Legs narrow, similar in length; femora compressed laterally; male with metafemur swollen basally, posterior margin curved; tibiae shorter than femora, weakly dilated to narrow apex. Protibia not dilated at apex, with fine setal fringe on ventral, lateral margin smooth without stout spinules; few small stout spinules at disto-lateral angle. Meso- and metatibia narrow, similar to protibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV lacking pair of erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left side of animal and resting on side (Fig. 8 G). Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat spatula-shaped parameres. Parameres with long setae apically; length / width ratio 2.00. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 5. Female unknown, but as with C. abyssa, they are expected to have narrow metafemora.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFCFF93FF70FB26FBD1C06A.taxon	materials_examined	Type locality. Vietnam, Vinh Phuc Province. Range. Vietnam, Vinh Phuc Province. Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ VIETNAM: Vinh Phuc Prov., Me Linh Biodiv. Sta.; Dai Lai Lake, 27 – 29. ix. 2013; 100 m, Col. J. B. Heppner ”; 2) [rectangular; red, printed in black ink] “ HOLOTYPE ♂ Cycadophila cyclochasma P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFCFF93FF70FB26FBD1C06A.taxon	etymology	Etymology. The species epithet comes from the Greek, ‘ cyclo’ - for circular, and ‘ khasma’ — for deep void, pit, in combination referring to the ‘ circular pit’ at the submental-gular juncture.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFCFF93FF70FB26FBD1C06A.taxon	discussion	Remarks. Cycadophila cyclochasma is most similar in all morphological characters to C. abyssa. The key differences are subtle, but these differences are consistent in available specimens. Over 15 species of Cycas occur in the general region of northern Vietnam and adjacent regions of southeastern China (Osborne et al. 2007). All these Cycas belong in the Section Stangerioides. Only one species of Cycas, C. balansae, belonging in Subsection Stangerioides, is currently known from the type locality of Vinh Phuc province and this species ranges north into Guangxi province, China. The closely related Cycadophila abyssa is known from Cycas hainanesis on Hainan Island and habitat of C. fairylakea in Guangdong province, China (both belonging to Section Stangerioides, Subsection Taiwaninae), farther to the east. Based on these differences in geographical distributions, available hosts and morphology we consider these beetles to be separate species. Although molecular evidence is not currently available to confirm this conclusion, describing the species officially recognizes these differences and provides a name for future research. Cycadophila cyclochasma and C. abyssa differ from all other members of the genus by sharing a number of unique features on the head, most notably the deep pit at the submental-gular juncture, but also the swollen head that creates a deep groove around the eyes.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFAFF91FF70F94DFD98C16C.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above, readily distinguished from other member of the genus by the narrowed body, nearly quadrate pronotum, and color pattern. Adult description. Length 4.06 – 4.29 mm, width 1.53 – 1.59 mm. Body in dorsal view elongate, sides nearly straight and parallel, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color dark reddish-brown, pronotum with orange-brown sides, elytra mostly orange-brown with dark maculation along suture nearly touching base, widening broad central spot, ending 1 / 3 elytral length from apex; dorsal surface distinctly punctate, surface appearing slightly shiny, dorsal punctation with short procumbent hairs, ventrally mostly covered with short procumbent setae. Head in dorsal view conical, gradually narrowed apically, surface convex, distinctly punctured; width 1.0 mm; dorsal interocular distance 0.50 – 0.58 mm, head width / dorsal interocular distance ratio 1.65 – 1.73, ventral interocular distance 0.38 – 0.40 mm, head width / ventral interocular distance ratio 2.38 – 2.53. Eye globular; bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 13: 11: 13: 10: 10: 10: 10: 10: 15: 15: 18; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palp palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow depression present with setose punctures.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFAFF91FF70F94DFD98C16C.taxon	description	Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.70 – 0.73, convex, nearly parallel-sided; anterior angles projecting, weakly angulate; lateral carinae nearly straight, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 o; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.16 – 0.18. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate. Hypomeron with coarse punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate nearly parallel sided, convex; length / width 3.40 – 3.63, greatest width near midlength; with marginal bead basally, scutellary striole with 8 – 10 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta length less than interpuncture distance. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; femora compressed laterally; tibiae shorter than femora, weakly dilated to narrow apex. Protibia not dilated at apex, with fine setal fringe on ventral, lateral margin smooth without stout spinules. Meso- and metatibia narrow, similar to protibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.5 0. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 2: 9. Female not notably different externally from male paratype.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFAFF91FF70F94DFD98C16C.taxon	materials_examined	Type locality. Tienmuschan, N. W. China. Range. China. The region cited in the label of the holotype, N. W. China, would correspond in modern China to arid Xinjiang Province and may be an error. The type locality, Tienmuschan, most closely corresponds with Tienmushan, Zhejiang Province in coastal eastern China, which borders the northern boundary of Fujian Province, the other known locality of this species. Material examined. Holotype female with the following labels (Fig. 9 D): 1) [rectangular; white; printed in black ink] “ Tienmuschan, N. W. China Rtt. ”; 2) [rectangular; red, printed and hand written in black ink] “ Holo- TYPUS discimaculata M. ”; 3) [rectangular; white; hand written in black ink] “ Pharaxanota [sic] discimaculata Mad. ”. Deposited in NHMB (examined). Paratype male with the following labels: 1) [rectangular; white; printed in black ink, with black border] “ Tienmuschan, N. W. China Rtt. ”; 2) [rectangular; pink, printed and hand written in black ink] “ Para- COTYPUS discimaculata M. ” (NHMB, examined). Others examined: CHINA: Fujian Prov., Wuyishan City, Guadun Vill.; 27 o 44 ' N, 117 o 33 ' E, 1200 – 1500 m; 25 - V- 2012, PENG & DAI leg. (2). Deposited in SNUC, FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFFAFF91FF70F94DFD98C16C.taxon	discussion	Remarks. Two species of Cycas are native to Fujian Province, China (Ye 1999; Liu & Qin 2004), one of the known localities for this beetle. Author W. Tang recently visited Wuyishan in Fujian, one of the collection localities and found Cycas cultivated in that city.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF9FFAFFF70FF7CFA81C14A.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above distinguished by the very narrowed legs, protibia lack of stout setae apically, short supraocular striae, and broad pronotum, width almost 2.0 0 times length. Adult description. Length 4.57 mm, width 1.76 mm. Body in dorsal view elongate, sides evenly arcuate toward ends, greatest width at middle of elytra; in lateral view convex dorsally. General body color dark brown; appendages, head and lateral pronotum reddish-brown; elytra dark brown with reddish brown mark on humerus and apical third; dorsal surface distinctly punctate, surface appearing shiny, dorsal punctation with short procumbent hairs, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF9FFAFFF70FF7CFA81C14A.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface convex, distinctly punctured; width 0.86 mm; dorsal interocular distance 0.53 mm, head width / dorsal interocular distance ratio 1.61, ventral interocular distance 0.40 mm, head width / ventral interocular distance ratio 2.18. Eye globular; bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 2 / 3 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 16: 13: 13: 10: 10: 10: 10: 10: 15: 15: 19; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow depression with setose punctures is present on each side of middle. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.60, convex, nearly parallel-sided; anterior angles projecting, weakly angulate; lateral carinae nearly straight, diverging slightly posteriorly, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 o; posterior margin straight laterally, weakly lobed medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.13. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate, weakly convex in lateral view, weakly projecting. Hypomeron with coarse punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate nearly parallel sided, length / width 3.31, greatest width near midlength; with marginal bead basally, scutellary striole with 10 punctures; intervals of striae with fine, distinct punctures. All elytral punctures bearing a single short seta; seta length less than interpuncture distance. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; femora compressed laterally, weakly widened medially; tibiae shorter than femora, narrow and parallel-sided to apex. Protibia not dilated at apex, with fine setal fringe on ventral, lateral margin smooth without stout spinules. Meso- and metatibia narrow, similar to protibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV apparently lacking erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male unknown, sexual dimorphism unknown.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF9FFAFFF70FF7CFA81C14A.taxon	materials_examined	Type locality. Thailand, Chieng Mai. Range. Thailand, Chieng Mai. Material examined. Holotype (by designation) female with the following labels: 1) [rectangular; white; printed in black ink] “ THAILAND: Chiengmai, 12 - 19 - V- 1958, Viniol Notananda ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♀ Cycadophila eurynota P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the USNM.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF9FFAFFF70FF7CFA81C14A.taxon	etymology	Etymology. The species epithet combines the Greek ‘ eurys’, meaning broad, with the shortened name for the structure pronotum, and Latinizing them into the feminine ‘ eurynota ’.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFF9FFAFFF70FF7CFA81C14A.taxon	discussion	Remarks. The color, head and pronotal characters place this species in the nigra species group; however, the shape of the pronotum is unique among members of this group. Other characters, like the narrowed legs, are found on few others in the genus. These unique characters warrant description of the species even without a male. In color pattern, C. eurynota fits the description of C. intermedia. However, Chûjô (1967) distinctly states the tibia are triangularly widened, where they are decidedly not so in C. eurynota. The type locality, Chieng Mai, or Chiang Mai Province, has three species of Cycas, C. pectinata, C. siamensis and C. simplicipinna (Tang et al. 1997; Tang, unpublished data). Surveys of the first two species have not yielded specimens matching C. eurynota. Wild coning plants of C. simplicipinna have not been adequately surveyed and Coleoptera visitors, if any, are not known.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC7FFADFF70FF7CFDBCC794.taxon	description	Head strongly and closely punctured; frons lightly convex, with a distinct depression at each side of the middle of the anterior part; clypeus not limited from frons, distinctly narrowed forwardly, gently rounded at the front border in a dorsal aspect. Eyes large, much longer than it is broad, distinctly convex and very finely facetted. Antennae robust; 1 st segment strongly thickened and dilated terminally, 2 nd segment as broad as it is long, 4 th ~ 7 th segments moniliform, 3 rd segment distinctly longer than wide, longer than either the 2 nd and 4 th segments, 8 th segment transverse and broader than the 7 th segment but far smaller than the 9 th segment, 9 th and 10 th segments strongly enlarged with the former one semicircular and the latter transverse, 11 th segment transverse subovate and distinctly wider than long, 11 th segment narrower than either the 10 th and 9 th segments but very much larger than the 8 th segment. Pronotum somewhat broader than long, slightly narrowed forwardly; front border nearly straight and immarginate at the median part, but narrowly concave and marginate at the lateral parts; four corners forming a right angle in each, but not sharply angulate at each tip; lateral borders distinctly marginate, nearly straight, but gently curved inwardly at each, the anterior and basal, extremity; basal angles each forming almost a right angle; basal border narrowly marginate and arched posteriorly at the median part; dorsally lightly convex from side to side, but rather strongly depressed at each laterio-anterior area, strongly and rather loosely punctured on the median area, but very finely and scantily punctured on each lateral area and impunctate on each laterio-marginal area; laterio-basal foveae deeply impressed, strongly elongate triangular in each shape and almost parallel with each other. Scutellum strongly transverse, finely angulate at each point touching the base of the sutural border of elytra, flat and very finely but very sparingly punctured on the surface. Elytra slightly wider at the basal extremity than at the pronotum, but broader at the post-humeral area, gently narrowed posteriorly at the posterior part and widely rounded at the apical border; dorsum lightly convex, narrowly but distinctly marginate at the basal border, with a row of distinct punctures closely along this border; each elytron with 11 files (including a short scutellar and an external marginal file) of distinct punctures (these files of punctures becoming finer and uneven at the posterior area of elytron), and each interstice of these files of punctures with a regularly or rather irregularly arranged file of very fine punctures (the size of these fine punctures consisted of two or three kinds); humeri not so markedly elevated, with a few files of rather larger and fine punctures; elytral epipleuron broad at the base and rather markedly narrowed posteriorly at the basal one-third, but continued to the apex of elytra in a similar width at the posterior two-thirds, finely but not so closely punctured on the whole surface. Underside closely, rather strongly and evenly pubescent-punctate, but the lateral areas of prothorax and the median area of metathorax very finely and scantily pubescent-punctate; abdomen strongly, closely and evenly pubescent-punctate. Legs rather robust, with the femora dilated medially, tibia dilated terminally and each 1 st tarsal segment not flattened. ”	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC7FFADFF70FF7CFDBCC794.taxon	materials_examined	Type locality. Laos. Type. According to Chûjô (1967), the holotype is a male with label data: “ LAOS: Muong Sing (Alt. 650 m), NW. of Luang Prabang. 6 - 10. VI. 1960, S. et L. W. QUATE leg. ”. Chûjô (1967) reported it to be deposited in the Bernice P. Bishop Museum Museum, Honolulu, Hawaii, where it is registered as number 11005. But, it is presently missing and not available for study (Sheperd Myers, pers. comm. 2012). It is the only known specimen of the species.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC7FFADFF70FF7CFDBCC794.taxon	discussion	Remarks. Chûjô’s description and key to C. intermedia are lengthy. Yet, they provide no usable characters that confirm this species is a member of Cycadophila. Other Coleoptera have been mistakenly placed in Pharaxonothinae and Pharaxonotha as pointed out by Skelley (2013). Until the type of C. intermedia is available for study, we can neither confirm nor refute the generic or familial placement of this species. Assuming C. intermedia is a member of Cycadophila, the only usable comments in the description are the pronotal shape, dark body, and elytral color pattern which is an intermediate form as discussed for C. yunnanensis, and a member of the nigra species complex. Considering our hypotheses that cryptic species remain to be discovered in Cycadophila, we prefer to consider C. intermedia as an incertae sedis, and not to synonymize it until more material has been studied from Laos. The locality Muong Sing in Laos is close to the border region of China, where populations of Cycas sp. aff. collina are known (see Remarks under Cycadophila collina). Although nine species of Cycas are recognized as being present in Laos by Averyanov et al. (2014), this locality is not identified as harboring cycad populations. It is possible that the beetle has no association with cycads.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC4FFACFF70FE14FA97C6EC.taxon	diagnosis	Adult diagnosis. A member of the C. (Cycadophila) nigra species group as discussed above distinguished from related species by its larger entirely brown body, weakly depressed submental-gular region, distinct supraocular stria that is half length of the eye, and male with large angulate swelling at the middle of the inner mesotibial margin. A full description and images are presented in Xu et al. (2015).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC4FFACFF70FE14FA97C6EC.taxon	materials_examined	Type locality. Kanara, S. India. Range. China, S. India, Thailand, Vietnam. Material examined. Lectotype and paralectotype of Thallis nigra Gorham, holotype of Pharaxonotha indica Grouvelle, and other materials as detailed in Xu et al. (2015). Others examined: CHINA, Yunnan Province, Red River Valley, nr. Man Hao, 23 ° 01 ' N, 103 ° 24 ' E, 955 m, 12 May 1996, W. Tang, ex near receptive ♀ strobilus of Cycas diannanensis (type locality), WT 50 (2); Guanping Reserve, 22 ° 10 ' N, 110 ° 50 ' E, 10 May 1996, W. Tang, ex spent ♂ strobilus of Cycas dolichopylla [Cycas sp. aff. collina], WT 44 (3); VIETNAM, Son La Prov., near Mai Son, 21 ° 08 ' N, 104 ° 11 ' E, 4 Apr. 1996, S. – L. Yang, ex ♂ strobilus of Cycas collina in monsoon forest, SLY 777 (19); Hoa Binh Prov., LacThuy Dstr., Phu Lao comm., Thuy Dstr., Phu Lao comm., 20 ° 33 ' 19 ″ N, 105 ° 45 ' 42 ″ E, 50 – 100 m, 25 - IV- 2015, N. S. Khang, ex Ƌ Cycas hoabinensis cone (NSK 795 b) (1). This additional material examined is deposited at ANIC and FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC4FFACFF70FE14FA97C6EC.taxon	discussion	Remarks. Adult specimens matching this species were found in small numbers in cones of one of two populations of Cycas debaoensis surveyed in China (Xu et al. 2015). These authors found no matching larvae in the cones and suggested that this species does not reproduce in Cycas cones as do some other species of Cycadophila. Their apparent distribution into southern India, well beyond the range of C. debaoensis or related species, suggest that this beetle is not a specialist on C. debaoensis, but an opportunistic visitor that may feed and reproduce on alternative hosts. It is unclear if specimens presently considered C. nigra represent one widespread species or a couple of regional species. The female types of C. nigra are from southern India, while other available materials are from northern India, southern China and northern Vietnam. Additional series of specimens from other areas are needed to better understand sexual dimorphisms, population variations, and molecular relationships to decide if only one species is present. For now, C. nigra is considered a single widespread species needing more research. Arrow (1925) pointed out that the types of Gorham’s T. nigra and Grouvelle’s P. indica are identically labeled and synonymized them. They are all females with quite similar morphology; Gorham’s specimens being larger and darker and Grouvelle’s specimen being smaller and teneral. These differences probably led to the early confusion and separate generic placement in the original descriptions. We have no doubt that they belong in Cycadophila; however, there is doubt in the species synonymy of P. indica with C. nigra. Our key characterizes P. nigra to be a larger species (> 4.00 mm), but the type of P. indica is only 3.00 mm. This smaller size and other subtle yet variable characters are like C. yunnanensis or C. vittata. It is well documented that more than one species can be found on the same cycad cone, thus locality of capture is not a valid reason to synonymize species. Unfortunately, the type of P. indica is a female and males are needed to confidently characterize the species. Until additional series of specimens are available from southern India to know the taxa that occur there, we can only guess the true identity of P. indica. For now, we leave it as a synonym of C. nigra, pending future work with additional materials.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC5FFAAFF70FEF1FAD0C334.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above, distinguished by the narrowed legs, protibia lacking stout setae apically, short supraocular stria, presence of a small temple behind eye, pronotum notably reflexed along entire lateral carina, and pronotal anterior margin straight. Adult description. Length 3.50 – 4.10 mm, width 1.38 – 1.40 mm. Body in dorsal view elongate, sides nearly straight and parallel, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color dark reddish-brown; dorsal surface distinctly punctate, surface appearing shiny, dorsal punctation with short procumbent hairs, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC5FFAAFF70FEF1FAD0C334.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface convex, distinctly punctured; width 0.72 – 0.81 mm; dorsal interocular distance 0.47 – 0.56 mm, head width / dorsal interocular distance ratio 1.46 – 1.55, ventral interocular distance 0.35 – 0.37 mm, head width / ventral interocular distance ratio 2.07 – 2.19. Eye globular, surrounded by distinct depression (groove), posteriorly bordered by distinct ridge (temple); bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 18: 11: 12: 10: 10: 10: 10: 10: 13: 13: 20; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with small lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow but distinct depression present on each side with setose punctures. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides except anterior margin; length / width ratio 0.69 – 0.70; disc convex medially, becoming reflexed and flattened laterally; anterior angles not projecting, rounded; lateral carinae nearly straight, nearly parallel, weakly converging posteriorly; lateral carinae with thin bead, bead bearing row of setose punctures; posterior angles rounded, nearly 90 °; posterior margin straight laterally, weakly lobed medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.19 – 0.20. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate and weakly convex in lateral view, weakly projecting. Hypomeron with coarse punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate nearly parallel sided, length / width 3.27 – 3.80, greatest width near midlength; with marginal bead basally, scutellary striole with 10 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta length approximately interpuncture distance. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; femora compressed laterally; tibiae about same length as femora, narrow and parallel-sided to apex. Protibia not dilated at apex, with fine setal fringe on ventral, lateral margin smooth without stout spinules. Meso- and metatibia narrow, similar to protibia. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV apparently lacking erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat spatula-shaped parameres. Parameres short, with long setae apically; length / width ratio 2.00. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 3; flagellum very fine, or at least not sclerotized. Female unknown.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC5FFAAFF70FEF1FAD0C334.taxon	materials_examined	Type locality. Yibin, Sichuan, China (USNM). Range. China (Sichuan), Thailand (Kaen Municipality). Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ Suifu, Oct. 1924, 1 – 3000 ft. ”; 2) [rectangular; white; printed in black ink] “ Szechuen, CHINA, DCGraham ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila torquata P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the USNM. Paratype male: THAILAND: Kaen Municipality, 9 - V- 1954, R. E. Elbel (1). Deposited in the FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC5FFAAFF70FEF1FAD0C334.taxon	etymology	Etymology. The species epithet, ‘ torquata ’, is Latin for adorned with a necklace or collar, in reference to the prominent ridges (temples) behind the eyes.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC5FFAAFF70FEF1FAD0C334.taxon	discussion	Remarks. The type locality Suifu, Szechwan, is now Yibin, Sichuan. The collector, David Crockett Graham, was a missionary studying Chinese language and culture. While performing his studies in Suifu, he made occasional expeditions in the region to collect specimens for the Smithsonian Institution. His field note books reside in the Smithsonian archives (Anonymous 2015). Unfortunately, they contain no entries surrounding the dates the holotype was collected, thus we have no further information on the specimens habits. The type locality, Yibin, is approximately 100 – 150 km north of the northernmost populations of wild Cycas, including Cycas panzhuaensis in Panzhihua, Sichuan and C. guizhouensis in Guizhou province. The other locality for this species, Kaen Municipality, is likely Kanchanaburi province, Thailand (see Remarks under Cycadophila convexa) where surveys on three species of Cycas yielded no specimens matching Cycadophila torquata. It is likely this species has no association with Cycas. The occipital ridge, lack of abdominal calli, head characters, and twisted male genitalia place C. torquata in the nigra group of Cycadophila. This is a strange beetle, looking quite different from the others by the pronotal shape and presence of a temple behind the eye. The latter character is only known from P. kirschii in the group of cycad inhabiting genera. Cycadophila torquata and C. convexa share a straight anterior pronotal margin which is not seen in any other member of Cycadophila, but differ notably in other pronotal characters. Cycadophila torquata and C. abyssa share the straight, parallel-sided lateral pronotal margins and groove around the eye. The holotype and paratype are similar in all diagnostic characters. The Chinese holotype differs in having deeper depressions at the submental-gular suture, and the temple behind the eye more prominent. Since neither of these specimens have any indication of a cycad association, we cannot assume one exists. Thus, we cannot assume that any population is isolated due to an association with a particular cycad species. Additional specimens and possibly molecular analysis are needed before we can consider these populations as different species.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC3FFA8FF70FAB4FCF2C334.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above, distinguished from other species by the narrow protibia lacking stout setae apically, short supraocular stria, coarser pronotal punctation, and having a large elytral maculation. Adult description. (Based on holotype) Length 4.00 mm, width 1.50 mm. Body in dorsal view elongate, sides arcuate, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid pale brown, elytra yellow-brown with dark maculation along suture from base near scutellum nearly to apex; dorsal surface distinctly punctate, surface shiny, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC3FFA8FF70FAB4FCF2C334.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; head width / dorsal interocular distance ratio 1.59, head width / ventral interocular distance ratio 2.10. Eye globular; bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 18: 12: 13: 10: 10: 10: 10: 10: 15: 15: 20; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 4: 3: 7. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a vague discolored depression is present marked by last set of submental punctures indicating remains of suture. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.62, convex, sides arcuate; anterior angles rounded; lateral carina with thin marginal bead, bead bearing row of minutely setose punctures; posterior angle exteriorly angulate; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.16. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process truncate and convex. Hypomeron with few punctures laterally, medially without longitudinal striations. Scutellar shield transverse, posterior margin rounded, somewhat pentagonal. Elytra elongate, evenly arcuate along sides, convex; length / width 3.30, greatest width near midlength; with marginal bead basally, scutellary striole with 10 punctures; intervals of striae with fine, distinct punctures. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; procoxae oval; mesocoxae globular; metacoxae transversely elongate-oval; trochanters obliquely truncate apically; femora compressed laterally; tibiae shorter than femora, narrow at apex. Protibia narrow, weakly angled on lateral apex, apex fringed with small, hair-like spinules, lateral margin lacking spinules. Meso- and metatibia narrowed, obliquely angled at lateral apex. Male metatibia with slight rounded swelling on inner apical half. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV apparently lacking erect sensory hairs located on either side of the middle; II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres short, with long setae apically; length / width ratio 2.00. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 5. Female unknown.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC3FFA8FF70FAB4FCF2C334.taxon	materials_examined	Type locality. Burma (now Myanmar). Range. Burma (now Myanmar). Material examined. Holotype male with the following labels (Fig. 12 F): 1) [rectangular; white; hand written in black ink] “ Maymyo VI, 16 H. L. Andrewes ”; 2) [rectangular; white with yellow horizontal line; printed in black ink] “ Maymyo, Burma, H. L. Andrewes ”; 3) [rectangular; white; printed in black ink] “ Andrewes Bequest B. M. 1922 – 221. ”; 4) [white circle, ringed with red ink; printed in black] “ Type ”; 5) [rectangular; white; hand written in black ink] “ Pharaxonotha vittata, Arrow type ”. Deposited in the BMNH (examined).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC3FFA8FF70FAB4FCF2C334.taxon	discussion	Remarks. Arrow (1925) discussed the subtle central inter-marginal swelling on the hind tibia of the type specimen as a possible sexual dimorphism. This dimorphism and other characters distinguish it from other species. However, this tibial character is also found in C. yunnanensis which has vittate forms in China. Cycadophila vittata and C. yunnanensis may represent a single species with two color forms or a species complex. For now, we consider C. vittata and C. yunnanensis to be distinct and will pursue further research on their relationship when additional materials from Myanmar are available. Two species of Cycas, C. pectinata and C. siamensis, are currently known from Myanmar (Lindström, unpub. data).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC1FFA7FF70FAB4FE2AC7B1.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) nigra species group as discussed above distinguished from related species by its smaller body with or without color pattern, weakly depressed submental-gular region, distinct supraocular stria less than 1 / 3 length of the eye, and male with small rounded swelling at the middle of the inner mesotibial margin. A full description and images are presented in Xu et al. (2015).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC1FFA7FF70FAB4FE2AC7B1.taxon	materials_examined	Type locality. Yunnan, China. Range. Known from Guangxi and Yunnan provinces, China, Manipur state, India and Laos. Material examined. Lectotype, 5 paralectotypes and other specimens as discussed in Xu et al. (2015). Others examined: CHINA, Yunnan Province, Guanping Reserve, 22 ° 10 ' N, 110 ° 50 ' E, 10 May 1996, W. Tang, ex spent ♂ strobilus of Cycas dolichopylla [Cycas sp. aff. collina], WT 44 (4); LAOS, Houphan Province, Viengxay Ds., Vieng Xai vill., 20 ° 23 ′ 41 ″ N, 104 ° 13 ′ 44 ″ E, 7 - IV- 2015, N. T. Hiep, L. Averyanov, N. S. Khang, N. Q. Hieu, T. Maisak, Somneux (7). These additional materials examined are deposited at ANIC and FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC1FFA7FF70FAB4FE2AC7B1.taxon	discussion	Remarks. Xu et al. (2015) detected large numbers of adults of this species in male cones of Cycas debaoensis that had finished shedding their pollen. However, no larvae of the species was detected, suggesting that this species is an opportunistic visitor that does not reproduce in male Cycas cones and that it may feed and reproduce in alternative hosts. DNA analysis of the 16 S rRNA gene suggests a complex of three cryptic species (see Fig. 1). Three color morphs can be discerned, but none are confined to any of the three haplotypes detected. Unfortunately, our morphological analyses have not yielded any other characters to support that more than one species is present. For now, C. yunnanensis is considered to be a variably colored species occurring in southern China, northern Laos and northeast India that needs more detailed work. See Remarks under C. vittata and the Remarks of C. yunnanensis in Xu et al. (2015).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA7FF70FE37FF35C242.taxon	diagnosis	Adult diagnosis. The Cycadophila (Cycadophila) papua species group is distinguished from other members of Cycadophila by the body ventrally reddish-brown, appendages reddish-brown, dorsally body mostly black with reddish-brown margins on head and pronotum and central stripe on each elytron; head dorsal surface flattened in profile; supraocular stria present, long, full length of eye; maxillary palpi long, almost 2 × longer than labial palpi; ventral interocular distance approximately 1 / 3 – 1 / 2 head width; gular-submental suture primarily flattened, medially with shallow U-shaped depression deeper on sided, with setose punctures; pronotal anterior angles rounded, surface near anterior angle convex; lateral margins evenly arched most of length, basally appearing evenly sinuate; pronotal lateral carina thin in lateral view, with single row of setose punctuation on marginal surface; prosternal process narrow (<1 / 2 width of profemur), appearing rounded at apex, apex depressed, in lateral view convex; protibia weakly dilated distally, lacking fringe of stout setae along apical and lateral margins; coxal line behind mesocoxae with weakly impressed large punctures; male genitalia with tegmen moderately twisted basally, penile strut about 4.0 0 × length of median lobe; and sexual dimorphism present with males having more slightly dilated pro- and mesotibial apices.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA7FF70FE37FF35C242.taxon	discussion	Remarks. The elongate maxillary palpi and broad flattened body readily distinguish the species of this group from all other. Their widespread distribution in the Philippines and New Guinea would indicate there are probably additional species in Indonesia occurring with the widespread and isolated members of Cycas (Lindstrom et al. 2009).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA5FF70FB25FC06C299.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Cycadophila) papua species group with long maxillary palpi as discussed for the group, recognized by its narrow orange mark on the elytra, prosternal process with coxal lines diverging posteriorly, and occurring in Papua New Guinea. Adult description. Length 6.85 – 6.90 mm, width 2.70 – 2.80 mm. Body in dorsal view elongate, sides evenly arcuate, greatest width at middle of elytra; in lateral view weakly convex dorsally. General body and appendage color solid reddish-brown, with nearly black base of head, pronotal disc, and sutural and lateral margins of elytra, remaining red-orange mark on elytra broad, at base from striae IV – VI; dorsal surface distinctly punctate, surface glossy, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA5FF70FB25FC06C299.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 1.10 – 1.17 mm; dorsal interocular distance 0.62 – 0.69 mm, head width / dorsal interocular distance ratio 1.70 – 1.77, ventral interocular distance 0.43 – 0.48 mm, head width / ventral interocular distance ratio 2.44 – 2.56. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 0.9 0 times pronotal width, and longer than head width, antennomere relative lengths from base to end approximately 16: 12: 14: 10: 10: 10: 10: 10: 14: 14: 18; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II and III longer than wide; IV elongate, apex densely papillate; relative lengths of II – IV approximately 4: 4: 8. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with small lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow U-shaped fovea is present, deepest laterally, bearing setose punctures. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.58 – 0.61, moderately convex, sides evenly arcuate entire length; anterior angles rounded; lateral carina with fine bead, bead bearing row of fine setose punctures; posterior angle exteriorly angulate; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.17 – 0.19. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex; coxal lines diverging posteriorly. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, evenly gradually arcuate laterally, convex; length / width 3.38 – 3.43, greatest width near midlength; with marginal bead basally, scutellary striole with 10 punctures, intervals of striae with distinct fine punctures. All elytral punctures bearing a single short seta; seta length barely reaching outside of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventral length. Legs narrow, similar in length; femora compressed laterally; tibiae shorter than femora, weakly dilated to narrow apex. Protibia weakly dilated at apex, apical and lateral margin lacking stout spinules, 2 – 3 small stout spines present on apical-lateral angle giving angle a sharper appearance. Meso- and metatibia less dilated, nearly parallel-sided at apex. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat spatula-shaped parameres with oblique apex. Parameres with long setae apically; length / width ratio 2: 1. Aedeagus with median lobe short, slender, highly sclerotized, apex acuminate. Length of median lobe to penile strut ratio is around 1: 4. Female similar to male; pro- and mesotibia narrower than male.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA5FF70FB25FC06C299.taxon	materials_examined	Type locality. Papua New Guinea, Nadzad. Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ Nadzab, Markam R. Val. NGuinea, Au, K. V. Krombein ”; 2) [rectangular; white; printed in black ink] “ ExColl KVKrombein ”; 3) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila papua P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the USNM. Allotype female, same data as holotype. Deposited in the USNM.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA5FF70FB25FC06C299.taxon	etymology	Etymology. Named for the region of New Guinea the types are from, Papua is used as a noun in apposition.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCEFFA5FF70FB25FC06C299.taxon	discussion	Remarks. Based on the fact that the closely related species, Cycadophila samara, in this species group, was collected on cycads, it is anticipated that C. papua also has a cycad association. The type locality, Nadzab, is situated in the highlands of New Guinea and is habitat for Cycas schumanniana (Hill 1994 a) and another species, C. rumphii, occurs along coastal habitats at lower elevations (Hill 1994 b).	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCCFFA3FF70FB02FD63C2FD.taxon	diagnosis	Adult diagnosis. A member of the Cycadophila (Cycadophila) papua species group with long maxillary palpi as discussed for the group, recognized by its broad orange mark on the elytra, prosternal process with coxal lines parallel, and occurring in the Philippines. Adult description. Length 5.90 – 5.95 mm, width 2.30 – 2.40 mm. Body in dorsal view elongate, sides evenly arcuate, greatest width at middle of elytra; in lateral view weakly convex dorsally. General body and appendage color solid reddish-brown, with nearly black base of head, pronotal disc, and sutural and lateral margins of elytra, remaining red-orange mark on elytra broad, at base from striae III – VIII; dorsal surface distinctly punctate, surface glossy, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCCFFA3FF70FB02FD63C2FD.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 0.95 – 0.98 mm; dorsal interocular distance 0.55 – 0.57 mm, head width / dorsal interocular distance ratio 1.72 – 1.73, ventral interocular distance 0.40 – 0.43 mm, head width / ventral interocular distance ratio 2.21 – 2.38. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 0.90 times pronotal width, and longer than head width, antennomere relative lengths from base to end approximately 17: 10: 12: 10: 10: 10: 10: 10: 12: 12: 16; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II and III longer than wide; IV elongate, apex densely papillate; relative lengths of II – IV approximately 4: 4: 8. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with small lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, exceptnear suture with submentum where a shallow U-shaped fovea is present, deepest laterally, bearing setose punctures. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.63 – 0.67, moderately convex, sides evenly arcuate entire length; anterior angles rounded; lateral carina with fine bead, bead bearing row of fine setose punctures; posterior angle exteriorly angulate; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.15 – 0.21. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex; coxal lines parallel. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, evenly gradually arcuate laterally, convex; length / width 3.25 – 3.57, greatest width near midlength; with marginal bead basally, striole with 10 punctures; intervals of striae with distinct fine punctures. All elytral punctures bearing a single short seta; seta length barely reaching outside of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventral length. Legs narrow, similar in length; femora compressed laterally; tibiae shorter than femora, weakly dilated to narrow apex. Protibia weakly dilated at apex, apical and lateral margin lacking stout spinules, 2 – 3 small stout spines present on apical-lateral angle giving angle a sharper appearance. Meso- and metatibia less dilated, nearly parallel-sided at apex. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat spatula-shaped parameres with oblique apex. Parameres with long setae apically; length / width ratio 2: 1. Aedeagus with median lobe short, slender, highly sclerotized, apex acuminate. Length of median lobe to penile strut ratio is around 2: 9. Female similar to male; pro- and mesotibia narrower than male.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCCFFA3FF70FB02FD63C2FD.taxon	materials_examined	Type locality. Philippines, Samar Island. Range. Philippines. Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ PHILIPPINES: Samar Is., N of Calbayog, 12 ° 5 ′ 6.4 ″, 124 ° 30 ′ 24.5 ″, ex young ♂ cone Cycas nitida, 13 - IV- 2011, T. Marler ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila samara P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA. Allotype and 3 Paratypes with same data as holotype. Deposited in the FSCA. Others examined: PHILIPPINES: Panay Island, Caticlan, 11 ° 52.368 ’ N 122 ° 01.927 ’ E, ex: Cycas edentata ♂ cone, 5 / 15 / 2010, T. Marler [specimen destroyed during DNA analysis; pin, label & photograph voucher remain].	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCCFFA3FF70FB02FD63C2FD.taxon	etymology	Etymology. The specific epithet is based on the island where the holotype was collected with the suffix ‘ - a’ added to make it euphonic with the genus name.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCCFFA3FF70FB02FD63C2FD.taxon	discussion	Remarks. A specimen yielding a DNA sequence of the 16 S rRNA gene that closely matches that of the type specimens was collected in Panay Island, Phillipines on Cycas edentata (see D 0081 in Fig. 1). The specimen was destroyed during DNA sampling. However, a photographic voucher was taken, which confirms a close morphological match with Cycadophila samara.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCAFFA0FF70FAFCFA8CC7B1.taxon	materials_examined	Type species. Cycadophila (Strobilophila) tansachai, n. sp., here designated.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCAFFA0FF70FAFCFA8CC7B1.taxon	diagnosis	Adult diagnosis. The subgenus C. (Strobilophila) is easily distinguished from other members of the genus by the pronotal lateral carina flat and thickened in lateral view with punctation scattered on carinal surface, by the presence of stout setae on the lateral margin of the protibia and sexual dimorphism, when present, on abdominal ventrite V, not legs as in other groups. Adult description. Length 3.60 – 6.40 mm, width 1.40 – 2.40 mm. Body: form elongate, not strongly parallel sided; dorsoventrally flattened to weakly convex; widest at middle of elytra; color reddish-brown without color pattern; dorsal punctation moderately coarse, evenly distributed, dense. Head: dorsal surface of head relatively flat; convex between eyes, clypeus flat apically; some with vague depressions near base of clypeus; clypeus truncate anteriorly; supraocular stria extending about 3 / 4 of length of eye from posterior end; transverse occipital ridge present; stridulatory file on occipital region of head present, one on each side, widely separated, parallel; eye prominent, coarsely faceted; lacking lateral temple behind eye. Mouthparts: mentum with large triangular base defined by carina; lateral pockets in front of carina small or indistinct; submentum and gula relatively flat, suture not obvious, medially with transverse, anteriorly concave depression with row of setose punctures. Pronotum: pronotal anterior marginal bead complete, fine; pronotal anterior margin near eyes emarginate, anterior angles rounded; surface near anterior angle convex; pronotal basal pits present, associated longitudinal groove present or not, groove up to 1 / 4 pronotal length; secondary transverse groove between pits present, broad, curving along posterior margin except medially where slightly separated from margin; pronotal lateral carina evenly arcuate, lacking modifications; pronotal lateral carinal bead thin in dorsal view; in lateral view thickening anteriorly, surface flattened laterally, appearing double edged. Scutellar shield broadly pentagonal. Elytra with coarse, dense interval punctures. Prosternum: pronotal hypomeron with longitudinal striations, weak in some. Legs: protibia dilated distally, angled laterally but without apical tooth; disto-ventral margin with fringe of stout ventrally directed, tooth-like spinules, some present on apical-lateral margin; meso- and metafemora elongate oval, meso and metatibiae moderately angularly dilated at apex. Abdomen: ventrites I – IV lacking pair of erect sensory hairs located on either side of the middle. Sexual dimorphism: not evident on legs. Male genitalia: median lobe and tegmen flattened laterally, weakly twisted basally; penile struts not fused, separate entire length, each 5.00 – 6. 0 0 × longer than median lobe; flagellum shorter than penile strut, not coiled; spiculum gastrale asymmetrical. Female genitalia: gonocoxites triangular, gradually narrowed posteriorly; apices of gonocoxites laterally with concave impression and setae. Gonostylus small, inserted at the concave impression of the gonocoxite, with several short setae and 1 long seta. Valvifers expanded posteriorly. Spermatheca elongated, C-shaped.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCAFFA0FF70FAFCFA8CC7B1.taxon	description	Larva. Description based on a larval morphotype found with adults of Cycadophila (Strobilophila) tansachai in Cycas elephantipes male cones. 16 S rRNA gene sequences from one individual of this larval morphotype from this host match those of adult C. (S.) tansachai, but a second sample of this larval morphotype from Cycas pectinata did not match any sampled adults of subgenus Strobilophila. This second larva may correspond with C. (S.) yangi adults, which have been collected together with adults of C. (S.) tansachai in Cycas pectinata. For further discussion, see paragraph prior to larval key and description of C. (S.) yangi below. Individuals up to 7.00 mm; body elongate, in dorsal view sides subparallel and narrowing anteriorly and posteriorly, cylindrical, lightly pigmented except for head and sections of tergum (Figs. 15 A – B). Head: sclerotized, dark brown with faint pattern of reticulation, smooth without ornamentation, protracted, prognathous and slightly flattened; epicranial stem absent, frontal arms lyriform and contiguous basally; median epicranial suture distinct, extending half the length of the frontal arms; 5 stemmata on each side; antenna length approximately half of midlength of head, 3 - segmented, antennomere I slightly shorter than wide, II more than 3 times as long as I, III slightly longer than I; frontoclypeal suture absent; labrum free. Mandibles symmetrical, incisor cleft with dorsal prong shorter than ventral prong, both prongs smooth; mola well-developed, asperate with asperites forming transverse rows; accessory ventral process present; prostheca hyaline, with single edge, broad basally, triangular with angulate apex (Fig. 15 F). Ventral mouthparts retracted, with well-developed maxillary articulating area; maxilla with transverse cardo, elongate stipes, 3 - segmented palp, terminal palpomere> 3 × its own width, mala falciform with apex divided into 3 lobes; labium more or less free to base of mentum, labial palps 2 - segmented, separated by 1.50 × width of basal palpomere (Fig. 15 G), distal palpomere length 2.00 × its own width. Hypostomal rods diverging; ventral epicranial ridges weakly developed. Thorax and abdomen: dorsal surface: anterior 1 / 2 of T 2 – 3 and A 1 – A 7, anterior 3 / 4 of A 8, central 1 / 2 of T 1 and all of A 9 pigmented dark brown, sclerotized; brown areas, except for anterior 1 / 4 – 1 / 3, more or less randomly covered with granules; sclerotized regions of T 1 – 3 and A 1 – 8 distinctly bisected by light-colored middorsal line; T 2 – 3 and A 1 – 8 with transverse row of 5 – 6 pairs of large tubercules along posterior margin and transverse row of 3 – 4 pairs of large tubercles 1 / 3 of length from anterior margin; tubercles appearing light colored and contrasting against dark background, setiferous; A 9 tergum with pair of urogomphi; urogomphi covered with granules basally, dorsal and lateral sides of base with 3 pairs of prominent setiferous tubercles, in dorsal view tubercles form approximately semicircular pattern, pair of tubercles closest to midline just anterior to urogomphi (labeled t 1 in Figs. 15 C – D) with hook at apex, base of this pair angled vertically, apical hook shaped like a bird head and neck with bill pointing posteriad (Figs. 15 A, C), top of bird head with a prominence with a single setae; bird beak length about 2.00 × its own vertical width, without sharply curved tip; urogomphi length from basal tubercle t 3 (see Fig. 15 D) to apex approximately 1.00 × length of remainder of segment, apical 1 / 3 of length smooth without tubercles. Legs 5 - segmented; coxae moderately widely separated, procoxae separated by> 1.00 × coxal width; meso- and metacoxae each separated by> 2.00 × coxal width (Fig. 15 E). Spiracles raised, annular-biforous.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCAFFA0FF70FAFCFA8CC7B1.taxon	etymology	Etymology. An intentional combination of the plant structure name ‘ strobili’, with the Greek - phil-, having a strong affinity or love. The name is feminine in gender.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFCAFFA0FF70FAFCFA8CC7B1.taxon	discussion	Remarks. Strobilophila is distinct among the cycad inhabiting erotylids by the strongly dilated protibia fringed with stout spinules apically and the thickened lateral pronotal carina. In size and superficial characters, Strobilophila resembles Xenocryptus tenebrionoides Arrow which inhabits cycads in Australia and has been cited as “ Xenocryptus ” in Tang et al. (1999). However, the genus Xenocryptus has internal abdominal calli, symmetrical male genitalia, and belongs in the Xenocryptinae (Leschen 2003). The distinction between the Pharaxonothinae and the Xenocryptinae is tenuous and in need of further study (Xu et al. 2015). This must be done considering all known taxa of these subfamilies, including other problematic and undescribed taxa that are cycad associates.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC9FFBEFF70FE37FBC5C084.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Strobilophila) as discussed above, more readily distinguished by a combination of characters than any single unique character, and by analysis of the 16 S rRNA mitochondrial gene. Cycadophila assamensis can be distinguished by the distally dilated protibia bearing narrowly spaced teeth along the apical third of the lateral margin, the distinct furrow on each side of pronotal base, elytral disc being mostly glabrous, the nearly straight basal pronotal margin, the lateral pronotal carina being abruptly curved inward at posterior angle pore which has a small acute projection behind it, and the terminal abdominal ventrite being sexually dimorphic (males with surface evenly coarsely punctate and setose; females with surface entirely setose and bearing a glossy area near the medial apex). Adult description. Length 4.69 – 5.19 mm; width 1.8 8 – 2.00 mm. Body in dorsal view elongate, sides nearly straight, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid reddish-brown [available specimens poorly preserved, darker; image of fresh specimen has color identical to C. tansachai]; dorsal surface distinctly punctate, surface appearing slightly dulled, dorsally appearing glabrous, with short procumbent hairs associated with punctation, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC9FFBEFF70FE37FBC5C084.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 1.00 – 1.10 mm; dorsal interocular distance 0.66 – 0.73 mm, head width / dorsal interocular distance ratio 1.50 – 1.52; ventral interocular distance 0.46 – 0.49 mm, head width / ventral interocular distance ratio 2.16 – 2.25. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 2 / 3 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 19: 13: 13: 10: 10: 10: 10: 10: 15: 15: 19; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 3: 3: 5. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.65 – 0.68, convex, evenly weakly arcuate most of length; anterior angles rounded; lateral carina with thickened bead, bead bearing scattered setose punctures; posterior angle exteriorly angulate, indented at pore; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.22 – 0.24. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, nearly parallel sided, convex; length / width 3.28 – 3.65, greatest width near midlength; with marginal bead basally, striae not impressed, scutellary striole with 10 – 12 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta visible but barely emerging out of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending about ¾ metaventrite length. Legs stout, similar in length; femora compressed laterally; tibiae shorter than femora, dilated to wide apex. Protibia triangularly dilated at apex, apex and apical 1 / 3 of lateral margin with stout spinules, marginal spinules separated by less than length. Meso- and metatibia less triangularly dilated. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight or weakly bisinuate (Figs. 16 F – G); ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae; ventrite V surface of male evenly coarsely and rugosely punctate, and entirely setose (similar to C. tansachai in Fig. 19 F), female evenly setose, basally with coarse punctures, with small area of fine punctures and glossy surface near medial apex (Fig. 16 G). Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.2 0. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 5. Female similar to male; except with pygidium truncate not as well developed; and dimorphic abdominal ventrite V surface as noted above.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC9FFBEFF70FE37FBC5C084.taxon	materials_examined	Type locality. India: Assam, Kamrup, North Guwahati. Range. India (Assam, Bihar). Material examined. Holotype (by designation) female with the following labels: 1) [rectangular; white; printed in black ink] “ INDIA: Assam, Kamrup, North Guwahati, ex ♂ cone Cycas pectinata, 2008, K. J. Singh ”; 2) [rectangular; pink; printed in bold black ink] “ PHOTO ”. 3) [rectangular; white; printed in black ink] “ collection date June-July 2007 ”; 4) [rectangular; white; hand written in black ink] “ INDIA: ASSAM, Kamrup, North Guwahati, Cycas pectinata, NG 1 ”; 5) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila assamensis P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA. Allotype male and 3 paratypes: same data as holotype (holotype & allotype, FSCA, 1); INDIA: Assam, Guwahati IIT Campus, ex ♂ cone Cycas pectinata, 2008, K. J. Singh (2). Deposited in the FSCA. Others examined: One additional adult male collected in association with larvae, currently considered this species: INDIA: Bihar, West Champaran, Valmiki Nat. Park, Mar 2015, K. J. Singh (1). Deposited in FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC9FFBEFF70FE37FBC5C084.taxon	etymology	Etymology. Named for the State where the type specimens were collected.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFC9FFBEFF70FE37FBC5C084.taxon	discussion	Remarks. A female holotype was chosen because it was the most intact specimen available for study and because it shows the sexual dimorphism used to distinguish the species. Cycadophila assamensis is most similar to C. tansachai, differing most notably in abdominal sexual dimorphism. Paucity of Indian materials prevents us from performing certain analyses and answering certain questions. Adult morphology allow us to distinguish the populations from India and propose a name for future research. Larvae from Bihar are also morphologically distinguishable from C. (Strobilophila) larvae collected in Thailand by the posteriad-tilting orientation of their t 1 tubercle (see Key to larvae above) vs. the vertically oriented t 1 turbercle in Thai specimens. While the Assam and Bihar populations are presented here as conspecific, we have doubts requiring further analyses. Thus, the type series is limited to the Assam population where we are able to make morphological comparisons with all known species. As noted above, molecular analyses failed with available material. The host for this species in Assam, Cycas pectinata, occurs at atypically low elevations for the species (52 – 192 m; see Singh & Singh 2014) vs. over 500 m for other localities of Cycas pectinata in the states of Manipur and West Bengal, India or in Thailand and provides evidence that this beetle is geographically isolated from other populations of the Strobilophila complex. See also remarks under C. kwaiensis.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD7FFBBFF70F8E1FC67C491.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Strobilophila) as discussed above, readily distinguished from the other member of the subgenus by the distally dilated protibia bearing narrowly spaced teeth along the apical third of the lateral margin, the distinct furrow on each side of pronotal base with basal margin weakly angulate at base of furrow, elytral disc being mostly glabrous, and the female abdominal ventrite V with distinct concavity at apex (Fig. 17 D). Adult description. Length 5.25 – 6.40 mm; width 2.00 – 2.40 mm. Body in dorsal view elongate, sides nearly straight, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid reddish-brown; dorsal surface distinctly punctate, surface appearing slightly dulled, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD7FFBBFF70F8E1FC67C491.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 0.83 – 1.00 mm; dorsal interocular distance 0.50 – 0.62 mm, head width / dorsal interocular distance ratio 1.61 – 1.66, ventral interocular distance 0.33 – 0.40 mm, head width / ventral interocular distance ratio 2.50 – 2.52. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 18: 13: 14: 10: 10: 10: 11: 11: 17: 17: 21; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II and III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 3: 3: 6. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.69 – 0.72, convex, nearly parallel-sided; anterior angles rounded; lateral carina with thickened bead, bead bearing scattered setose punctures; posterior angle exteriorly angulate, indented at pore; posterior margin projecting medially, margin weakly angulate at base of discal furrow; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.21 – 0.27. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, nearly parallel sided, convex; length / width 3.33 – 3.63, greatest width near midlength; with marginal bead basally, scutellary striole with 10 – 13 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta barely emerging out of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, laterally convex, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout, similar in length; femora compressed laterally; tibiae shorter than femora, dilated to wide apex. Protibia triangularly dilated at apex, apex and apical 1 / 3 of lateral margin with stout spinules, marginal spinules separated by less than length. Meso- and metatibia less triangularly dilated. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex (male) or apical concavity (female), apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 1: 2.2. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 2: 7. Extruded endophallus similar to that of C. (S.) tansachai (see Figs. 19 H – I), except with a cluster of spicules located beneath the basal hemisphere and spicules scattered along the central tubular portion. Female similar to male; with apical concavity on last abdominal ventrite.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD7FFBBFF70F8E1FC67C491.taxon	materials_examined	Type locality. Vietnam: Khanh Hao, Tan Lap village. Range. Vietnam (Binh Thuan, Khanh Hao). Material examined. Holotype (by designation) female with the following labels: 1) [rectangular; white; printed in black ink] “ VIETNAM: Khanh Hao, Tan Lap vill., 140 m, 11 ° 58 ′ 27 ″ N, 108 ° 03 ′ 47 ″ E, ex ♂ cone Cycas elongata, 5 - VIII- 2002, W. Tang & H. T. Nguyen ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♀ Cycadophila hiepi P. Skelley, G. Xu & W. Tang 2 0 1 7 ”. Deposited in the FSCA. Allotype and 69 paratypes: VIETNAM: [Binh Thuan] 2 km S. of Ca Na, 11 ° 20 ' N, 108 ° 51 ' E, 23 Feb. 1995, S. - L. Yang, ex ♂ strobilus of Cycas pachypoda in dry rocky open forest near sea, SYL 736 (1); Khanh Hao, Tan Lap vill., 140 m, 11 ° 58 ' 27 ″ N, 108 ° 03 ' 47 ″ E, ex ♂ cone Cycas elongata, 5 - VIII- 2002, W. Tang & H. T. Nguyen (Allotype, FSCA, 54); 15 km N of Nha Trang, 12 ° 18 ' N, 109 ° 11 ' E, 66 m, 20 Feb. 1995, S. - L. Yang, ex ♀ strobilus of Cycas elongata on limestone hills, SYL 721 (9); SYL 722 (3); SYL 723 (2). Paratypes deposited at ANIC, FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD7FFBBFF70F8E1FC67C491.taxon	etymology	Etymology. In honor of Mr. Hiep Tien Nguyen, of the Center for Plant Conservation in Vietnam, for his assistance in the field and as co-collector of type specimens of this species.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD7FFBBFF70F8E1FC67C491.taxon	discussion	Remarks. Unlike most other species we describe, C. hiepi has a female for a holotype because the female dimorphism is distinctive and unique. This species appears to be restricted to southern coastal Vietnam in Cycas elongata and C. pachypoda. Without locality data, the males are difficult to distinguish from C. tansachai, which occurs on Cycas clivicola lutea, C. elephantipes, C. pectinata and C. tansachana in Thailand. Some of these hosts also range into India, Myanmar, Cambodia and southernmost Vietnam.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD2FFB9FF70FD17FEB2C53E.taxon	diagnosis	Adult diagnosis. A member of the Cycadophila (Strobilophila) tansachai complex distinguished by the pronotal posterior lateral carina lacking an abrupt inward curve at posterior angle, lacking a distinct projection behind the posterior angle pore, a nearly straight basal pronotal margin, and with no dimorphism on the female abdomen. They are also distinguishable by DNA analysis of the 16 S rRNA mitochondrial gene (GenBank Accession number KY 365221). Adult description. Length 4.07 – 4.65 mm; width 1.58 – 1.70 mm. Body in dorsal view elongate, sides nearly straight, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid reddish-brown; dorsal surface distinctly punctate, surface appearing slightly dulled, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD2FFB9FF70FD17FEB2C53E.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 0.9 – 1.0 mm; dorsal interocular distance 0.60 – 0.66 mm, head width / dorsal interocular distance ratio 1.49 – 1.52, ventral interocular distance 0.38 – 0.43 mm, head width / ventral interocular distance ratio 2.34 – 2.47. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 21: 12: 11: 10: 10: 10: 10: 10: 15: 15: 19; antennomere I (scape) fairly large, slightly elongate, antennomere II – III slightly longer than IV – VIII; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 3: 3: 5. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.69 – 0.75, convex, nearly parallel-sided; anterior angles rounded; lateral carina with thickened bead, bead bearing scattered setose punctures; posterior angle exteriorly angulate, not indented at pore; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.19 – 0.23. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, nearly parallel sided, convex; length / width 3.50 – 3.67, greatest width near midlength; with marginal bead basally, scutellary striole with 10 – 13 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta barely emerging out of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout, similar in length; femora compressed laterally; tibiae shorter than femora, dilated to wide apex. Protibia triangularly dilated at apex, apex and apical 1 / 3 of lateral margin with stout spinules, marginal spinules separated by less than length. Meso- and metatibia less triangularly dilated. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.20. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 5. Female similar to male, except with pygidium truncate, not as well developed.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD2FFB9FF70FD17FEB2C53E.taxon	materials_examined	Type locality. Thailand: Kanchanaburi Province, 14 ° 2 ′ N, 99 ° 29 ′ E. Range. Thailand (Kanchanaburi). Material examined. Holotype (by designation, with damaged left elytron) male with the following labels: 1) [rectangular; white; printed in black ink] “ THAILAND, Kanchanaburi, 14 ° 2 ′ N, 99 ° 29 ′ E, karst tower, ex crown hair of Cycas sp. aff. pectinata, 29 - VII- 2002, W. Tang ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila kwaiensis, W. Tang, G. Xu & P. Skelley 2015 ”. Deposited in the FSCA. Allotype and 4 paratypes (same label data) deposited at FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD2FFB9FF70FD17FEB2C53E.taxon	etymology	Etymology. Named for the locality of collection in the vicinity of the Kwai River Bridge in Kanchanaburi Province, Thailand.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD2FFB9FF70FD17FEB2C53E.taxon	discussion	Remarks. This species of Cycadophila appears to be restricted to a currently undescribed species of Cycas that inhabits karst towers in Kanchanaburi Province at elevations below 100 m. Although Cycadophila tansachai occurs on Cycas pectinata some 100 km north at elevations above 600 m in mountains in Kanchanaburi Province, DNA analysis indicates that Cycadophila kwaiensis is also closely related to Cycadophila hiepi in Vietnam, over 1000 km to the east. This suggests that C. kwaiensis and its host cycad are relicts with formerly wider distribution in lowland forests.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD0FFB6FF70FCA7FD46C0AE.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Strobilophila) as discussed above, most readily distinguished by a combination of characters than any single unique character, and by analysis of the 16 S rRNA mitochondrial gene (GenBank Accession numbers KY 356222 – KY 365225, KY 356244 – KY 365245). Cycadophila tansachai can be distinguished by the distally dilated protibia bearing narrowly spaced teeth along the apical third of the lateral margin, the distinct furrow on each side of pronotal base, elytral disc being mostly glabrous, the nearly straight basal pronotal margin, the lateral pronotal carina being abruptly curved inward at posterior angle pore which has a small acute projection behind it, and abdominal ventrite V being sexually dimorphic (males with surface evenly coarsely punctate and setose; females with a glossy, glabrous area near the medial apex). Adult description. Length 4.06 – 4.53 mm; width 1.6 5 – 1.76 mm. Body in dorsal view elongate, sides nearly straight, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid reddish-brown; dorsal surface distinctly punctate, surface appearing slightly dulled, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and along lateral sides and declivity of elytra, ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD0FFB6FF70FCA7FD46C0AE.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 1.0 0 – 1.1 0 mm; dorsal interocular distance 0.63 – 0.73 mm, head width / dorsal interocular distance ratio 1.45 – 1.56, ventral interocular distance 0.45 – 0.50 mm, head width / ventral interocular distance ratio 2.10 – 2.17. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 18: 12: 12: 10: 10: 10: 10: 10: 17: 18: 19; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 3: 3: 5. Labium with mentum pentagonal with carina forming triangular plate projecting medially, lacking distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.66 – 0.68, convex, nearly parallel-sided; anterior angles rounded; lateral carina with thickened bead, bead bearing scattered setose punctures; posterior angle exteriorly angulate, indented at pore; posterior margin projecting medially; with narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.13 – 0.14. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, nearly parallel sided, convex; length / width 3.50 – 3.67, greatest width near midlength; with marginal bead basally, striae weakly impressed, scutellary striole with 10 – 13 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta barely emerging out of puncture. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout, similar in length; femora compressed laterally; tibiae shorter than femora, dilated to wide apex. Protibia triangularly dilated at apex, apex and apical 1 / 3 of lateral margin with stout spinules, marginal spinules separated by less than length. Meso- and metatibia less triangularly dilated. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each with pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae; ventrite V surface of male evenly coarsely and rugosely punctate, and entirely setose, female basally with coarse setose punctures, and small glossy glabrous area of fine punctures near medial apex (Figs. 19 F – G). Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.2 0. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 5. Endophallus, when inflated and extruded, basally hemispherical on dorsal side, with distal section of hemisphere covered with spicules (Figs. 19 H – I); central portion of extruded endophallus appearing tubular, membranous and without spicules or asperities; apical portion (not fully extruded in Figs. 19 H – I) appearing membranous and covered with asperities. Female similar to male, except with pygidium truncate, not as well developed; and dimorphic abdominal ventrite V surface as noted above.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD0FFB6FF70FCA7FD46C0AE.taxon	materials_examined	Type locality. Thailand: Chaiyaphum, Sai Thong Nat. Park, 15 ° 53.835 ' N, 101 ° 27.260 ' E. Range. Thailand (Chiang Mai, Chaiyaphum; Kanchanaburi, Sa Kaeo, Sariburi). Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ THAILAND: Chaiyaphum, Sai Thong Nat. Park, N 15 ° 53.835, E 101 ° 27.260, 862 m, ex ♂ cone Cycas elephantipes hybrid, 2 - II- 2011, W. Tang, A. Lindstrom et al. ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila tansachai P. Skelley, G. Xu & W. Tang 2 0 1 7 ”. Deposited in the FSCA. Allotype (FSCA) and 178 paratypes: THAILAND: [Chiang Mai] SW of Fang, 19 ° 38 ' N, 99 ° 07 ' E, 640 m, 7 Feb. 1995, W. Tang, S. - L. Yang, P. Vatcharakorn, ex pollen-shedding ♂ strobilus of Cycas pectinata in disturbed forest on limestone soil, SLY 690 (4); Chaiyaphum, Sai Thong Nat. Park, 15 ° 53.835 N, 101 ° 27.260 E, 862 m, ex ♂ cone Cycas elephantipes hybrid, 2 - II- 2011, W. Tang, A. Lindstrom et al. (holotype & allotype, FSCA, 124), Pa Hin Ngam Nat. Park, dry forest along ridge, 15 ° 38.732 N, 101 ° 23.483 E, 850 m, 2 - II- 2011, W. Tang, A. Lindstrom et al. (17); 15 ° 46 ' N, 101 ° 55 ' E, 30 May 1996, S. – L. Yang, ex ♂ strobilus of Cycas elephantipes, SLY 810 (3); Kanchanaburi Prov., near Thon Pha Phum, 14 ° 53 ' N, 98 ° 49 ' E, 700 m, 11 Feb. 1995, W. Tang, S. - L. Yang, P. Vatcharakorn, ex pollen-shedding ♂ strobilus of Cycas pectinata in disturbed forest, SLY 706 (9); SLY 707 (5); Loei Prov., Wat Pha Sawan, Karst cliff, 17 ° 07.15 N, 101 ° 56.00 E, 300 m, ex ♂ cone Cycas petraea, 3 - II- 2011, W. Tang, A. Lindstrom, A. Vogel et al. (5); [Sa Kaeo] near Aranyaprathet, 13 ° 22 ' N, 102 ° 18 ' E, 370 m, 25 Apr. 1996, W. Tang, ex ♂ strobilus of Cycas clivicola lutea in limestone cliffs, WT 21 (5); WT 22 (4); [Sariburi] near Sariburi, 14 ° 41 ' N, 100 ° 47 ' E, 370 m, 12 Feb. 1995, W. Tang, S. - L. Yang, P. Vatcharakorn, ex pollen-shedding ♂ strobilus of Cycas tansachana on limestone mountains in seasonally dry vegetation, SYL 715 (7). Paratypes deposited at ANIC, BMNH, FSCA, IZCAS, MNHN, NHMB, NZAC, USNM.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD0FFB6FF70FCA7FD46C0AE.taxon	etymology	Etymology. In honor of Mr. Kampon Tansacha, Director of Nong Nooch Tropical Botanical Garden in Thailand for his logistical and financial support of cycad beetle research.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFD0FFB6FF70FCA7FD46C0AE.taxon	discussion	Remarks. The detection of both adults and larvae together in cones of populations of Cycas petrea indicates that this is one of at least three species of Cycadophila that feeds and reproduces in Cycas cones. The two other species with DNA evidence indicating that larvae feed and develop in cones are Cycadophila debonica and C. fupingensis (Xu et al. 2015). The host for this species, the Cycas pectinata species complex, is widely distributed from Vietnam westward to Bihar State in India. The Cycas pectinata species complex includes C. elephantipes, C. pectinata, C. petraea, C. tansachana and C. clivicola lutea in Thailand (Yang & Meerow 1996; Lindstrom & Hill 2002). Geographical range and morphological variation of this host complex in India (Singh & Singh 2014) suggests that there may be recent speciation in the Cycadophila tansachai complex in that region, as supported by the similarities between it and C. assamensis.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFDFFFB3FF70F907FDA2C7E9.taxon	diagnosis	Adult diagnosis. A member of Cycadophila (Strobilophila) as discussed above, readily distinguished from the other members of the subgenus by the narrowed protibia bearing widely spaced teeth along the apical half of the lateral margin, lacking or having a greatly reduced furrow on each side of the pronotal base, entire elytral disc with short setae, and lacking notable sexual dimorphism. Adult description. Length 3.60 – 4.40 mm; width 1.4 0 – 1.68 mm. Body in dorsal view elongate, sides nearly straight, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color solid reddish-brown; dorsal surface distinctly punctate, surface appearing slightly dulled, dorsally appearing glabrous, with short procumbent hairs associated with punctation behind eyes and covering surface and declivity of elytra (best seen in profile), ventrally mostly covered with short procumbent setae.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFDFFFB3FF70F907FDA2C7E9.taxon	description	Head in dorsal view conical, gradually narrowed apically, surface flat to slightly convex, distinctly punctured; width 0.71 – 0.88 mm; dorsal interocular distance 0.45 – 0.59 mm, head width / dorsal interocular distance ratio 1.49 – 1.58, ventral interocular distance 0.38 – 0.47 mm, head width / ventral interocular distance ratio 1.86 – 1.87. Eye globular; bordered dorsally with supraocular stria; projected laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with short, dark, subcuticular suture on each side, approximately 1 / 4 width of frons in length, extending obliquely to lateral margin above and anterior to antennal insertion. Antennal length about 3 / 4 pronotal width, and slightly longer than head width, antennomere relative lengths from base to end approximately 19: 14: 13: 10: 10: 10: 10: 11: 15: 15: 16; antennomere I (scape) fairly large, slightly elongate, antennomere II slightly smaller than III; IV – VIII small, width equals length; club fairly large, IX – XI similar in length, XI globular. Clypeus truncate anteriorly, anterior margin with long setae; densely punctate; somewhat emarginate, with narrow margin. Maxillary palpomeres II, III slightly longer than wide; IV elongate oval, apex densely papillate; relative lengths of II – IV approximately 3: 3: 5. Labium with mentum pentagonal with carina forming triangular plate projecting medially, with small distinct lateral pockets; labial palpomere III elongate-oval. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum where a shallow transversely elongate fovea is present bearing dense setose punctures, fovea dark brown. Thorax with pronotum transverse in dorsal view, with marginal beads on all sides; length / width ratio 0.65 – 0.66, convex, nearly parallel-sided; anterior angles rounded; lateral carina with thickened bead, bead bearing scattered setose punctures; posterior angle exteriorly angulate, indented at pore; posterior margin projecting medially; with or without an indistinct short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, when present length of furrows relative to pronotal length ± 0.13. Prosternum with anterior margin slightly emarginate, finely denticulate with fringe of long, anteriorly directed setae; posterior prosternal process round and convex. Hypomeron with coarse punctures laterally, medially with longitudinal striations. Scutellar shield transverse, posterior margin projected, pentagonal. Elytra elongate, nearly parallel sided, convex; length / width 3.23 – 3.24, greatest width near midlength; with marginal bead basally, scutellary striole with 7 – 9 punctures; intervals of striae with distinct punctures. All elytral punctures bearing a single short seta; seta length less than interpuncture distance. Punctation on meso- and metaventrite distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs stout, similar in length; femora compressed laterally; tibiae shorter than femora, gradually dilated to narrow apex. Protibia narrowly dilated at apex, apex and apical 1 / 2 of lateral margin with stout spinules, marginal spinules separated by distance equal to their length. Meso- and metatibia slightly less dilated distally. Abdomen with 5 ventrites bearing fine punctures; anterior margin with intercoxal process narrow, with triangular point anteromedially, lateral edges slightly projected, lateral and posterior margins arcuate, converging posteriorly; anterior and posterior margins of ventrites more or less straight; ventrite I lacking subcoxal line, slightly longer medially than II; I – IV each possibly lacking a pair short hairs located adjacent to midline (often abraded); II – IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex bearing row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 ° toward the left side of animal and resting on side. Tegmen sclerotized, triangular, gently twisted; with anterior region ring-like, posterior region sheath-like, posterior margin slightly arcuate; lateral margins gradually converging posteriorly; dorsally with 2 large, elongate, somewhat coniform parameres. Parameres with long setae apically; length / width ratio 2.20. Aedeagus with median lobe elongate, slender, highly sclerotized, apex long and acuminate. Ratio of median lobe length to penile strut length 1: 3. Female similar to male, except with pygidium truncate, not as well developed.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFDFFFB3FF70F907FDA2C7E9.taxon	materials_examined	Type locality. Thailand: Chaiyaphum, Sai Thong Nat. Park, 15 ° 53.835 ' N, 101 ° 27.260 ' E. Range. Thailand (Chaiyaphum, Chiang Mai, Kanchanaburi, Prachuap Khiri Khan). Material examined. Holotype (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] “ THAILAND, [Prachuap Khiri Khan] Pran Buri near Kao Sam Roi Yot N. P., 12 ° 07 ' N, 99 ° 57 ' E, 23 Apr. 1996, W. Tang, ex ♂ strobilus of Cycas pranburiensis (WT 18) ”; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Cycadophila yangi P. Skelley, G. Xu & W. Tang 2017 ”. Deposited in the FSCA. Allotype and 11 paratypes: THAILAND: [Chiang Mai] SW of Fang, 19 ° 38 ' N, 99 ° 07 ' E, 640 m, 7 Feb. 1995, W. Tang, S. - L. Yang, P. Vatcharakorn, ex pollen-shedding ♂ strobilus of Cycas pectinata in disturbed forest on limestone soil, SLY 690 (4); Kanchanaburi Prov., near Thon Pha Phum, 14 ° 53 ' N, 98 ° 49 ' E, 700 m, 11 Feb. 1995, W. Tang, S. - L. Yang, P. Vatcharakorn, ex pollen-shedding ♂ strobilus of Cycas pectinata in disturbed forest, SLY 707 (1); [Prachuap Khiri Khan] Pran Buri near Kao Sam Roi Yot N. P., 12 ° 07 ' N, 99 ° 57 ' E, 23 Apr. 1996, W. Tang, ex ♂ strobilus of Cycas pranburiensis (WT 18) (holotype, FSCA; allotype, ANIC; 6). Paratypes deposited at ANIC, FSCA.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFDFFFB3FF70F907FDA2C7E9.taxon	etymology	Etymology. In honor of George (Si-Lin) Yang, founder and president of Seq-Gen, co-collector of type specimens of this species and many other specimens used in this study and for his financial support of this research.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
03A38781FFDFFFB3FF70F907FDA2C7E9.taxon	discussion	Remarks. Adults of Cycadophila (S.) yangi were collected together with those of C. (S.) tansachai in cones of Cycas pectinata in Thailand indicating that adults of these two species of subgenus Strobilophila may coinhabit a single male cone. A larval morphotype has been identified from these cones and the DNA of the 16 S rRNA gene was analyzed for two individuals of this larval morphotype. The DNA sequence of one larval individual matched that of adult C. (S.) tansachai. The other larva exhibited a DNA sequence substantially different from that of adults of three other Strobilophila species examined (see sample D 0333 in Fig. 1). Preservation of adult C. (S.) yangi available for study was insufficient for DNA analysis so we cannot match this unknown larva to C. (S.) yangi, however, it is likely the unmatched larva is C. (S.) yangi. Xu et al. (2015) demonstrated that adults of up to four species of subgenus Cycadophila and up to two species of larva may coinhabit a single Cycas cone. Similarly, adults of two species of subgenus Strobilophila may inhabit the same cones and two species are feeding and producing larvae within the same cones. There is no evidence that a third species of Strobilophila may be involved. Our current DNA analysis suggests that C. (S.) yangi may not belong within Strobilophila, but may be better placed in a separate clade. However, until stronger genetic evidence is available, we rely on morphological characters in assigning this species within Strobilophila.	en	Skelley, Paul, Xu, Guang, Tang, William, Lindström, Anders J., Marler, Thomas, Khuraijam, Jibankumar Singh, Singh, Rita, Rich, Stephen (2017): Review of Cycadophila Xu, Tang & Skelley (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas (Cycadaceae) in Asia, with descriptions of a new subgenus and thirteen new species. Zootaxa 4267 (1): 1-63, DOI: 10.5281/zenodo.575641
