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Tang, William, Dominguez, Hector Gomez (2022): Two new species of Pharaxonothinae beetles (Coleoptera: Erotylidae) inhabiting cones of the cycad Ceratozamia santillanii Pérez-Farr. & Vovides (Cycadales: Zamiaceae) in Mexico. Zootaxa 5150 (3): 428-442, DOI: https://doi.org/10.11646/zootaxa.5150.3.7

Pharaxonotha perezi Tang & Gomez Dominguez, new species (Figures 6A–K) Adult diagnosis. Pharaxonotha perezi can be distinguished from other described members of the genus by a combination of characters: Body length 2.59–3.51 mm; pronotum with anterior angles rounded, pronotal length relatively short with length/width (PL/PW) = 0.68–0.74; spermatheca compact with spermathecal and glandular duct adjacent and inserted in a common shallow pit (Fig. 6J); with known distribution in Mexico, state of Chiapas on Ceratozamia santillanii. Adult male description. Length 2.80–3.11 mm, width 1.10–1.24 mm (n = 6). Body in dorsal view elongateoval, greatest width at middle of elytra; in lateral view convex dorsally (Figs. 6A–C). Body color entirely brown; dorsal surface punctate, shining and appearing glabrous, short procumbent hairs associated with punctation on pronotum and elytra visible in profile only under high magnification, ventrally shining and covered with short procumbent setae. Head not broad, width = 0.70–0.73× pronotal width (Figs. 6D–F); in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, finely, moderately punctured, average distance between closest punctures 1–2× width of puncture; head width 0.62–0.69 mm; dorsal interocular distance 0.39–0.43 mm, head width/dorsal interocular distance ratio 1.57–1.64, ventral interocular distance 0.28–0.31 mm, head width/ventral interocular distance ratio 2.06–2.38. Eye with large black facets, about 3× diameter of head punctures. Antennal length slightly shorter than pronotal width, 1.3× head width; antennomere I (scape) fairly large, slightly elongate; antennomere II slightly shorter than III; IV–VIII small, width equals length; club fairly large, IX and X similar in length; XI enlarged, 1.6× longer than X, globular with rounded apex. Clypeus weakly concave anteriorly, moderately punctate. Mentum and submentum finely punctate, distance between nearest punctures approximately 2× own diameter, each puncture with a short seta (Fig. 6F). Gula area smooth, without punctation or setae, border with submentum usually with a shallow transverse depression. Thorax with pronotum transversely quadrate in dorsal view, length/width ratio 0.70–0.74; with distinct marginal beads laterally and basally, anteriorly with fine marginal bead laterally, but fading medially; convex; anterior angles broadly rounded, not projecting; posterior angles weakly developed, with small denticle at angle; lateral carinae parallel-sided or evenly shallowly arcuate for length; posterior margin slightly projecting medially, projection beginning approximately by pair of small, dark pores in margin located ¼ width from posterior angles, each pore marks base of a distinct sulcus extending anteriorly onto disc ¼ length of pronotum. Prosternum in ventral view convex, with few moderate punctations, distance between nearest punctures approximately 2× own diameter; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae, longest setae approximately 1/4 length of eye; prosternal process expanded apically, rounded and convex at apex. Hypomeron laterally with few minute punctures, medially lacking distinct longitudinal striations. Scutellar shield distinctly transverse, posterior margin weakly roundly pentagonal. Elytra in dorsal view elongate-oval, convex; length/width ratio1.67–1.74, greatest width near midlength; with distinct marginal line basally; 10 complete striae of moderate puncture size; scutellary striole extending ¼ elytral length, with 9–13 punctures; punctures of elytral striae as large as pronotal punctures, weakly impressed; intervals of striae with fine, shallow punctures; all punctures of elytra bearing a single short seta; seta only visible in profile, extending slightly out of puncture. Mesoventrite with strong punctation, distance between nearest punctures approximately equal to diameter of punctures, puncture depth moderate. Metaventrite glossy, with moderate punctation separated by 2–3× own diameter; finely punctured; entire surface convex, metathoracic discrimen extending approximately 3/4 metaventrite length. Legs narrow, relatively similar in length and shape. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora robust, moderately compressed laterally; tibiae shorter than femora, gradually dilated to obliquely truncate apices; protibia with apical lateral tooth distinct, with apical fringe of short spinules of concave ventral apical margin usually lacking near lateral tooth; meso- and metatibia with apical fringe of short spinules on anterior margin, finer setae on posterior margins. Abdomen. Ventrite I 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 longer medially than II; II–IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex; apical margin bearing short, sparse setae; all ventrites bearing moderate, shallow punctation across surface, distance to nearest puncture approximately 2× diameter of puncture, punctures bearing mostly reclining setae; ventrite V with setae length nearly uniformly approximately 2× diameter of puncture; I–IV each with 2 or more median pairs of longer, semi-erect sensory hairs (difficult to see in poor lighting, often abraded). Male genitalia (n=2) with elongate, slightly flattened cylindrical median lobe, long coiled flagellum and dorsoventrally flattened tegmen with basal piece ring-like, similar to all others in the genus; as in all other known members of the genus, tegmen and median lobe held upside down while in repose inside body (Tang et al. 2020) (Figs. 6G–H). Parameres of tegmen in dorso-ventral view nearly parallel-sided, expanding slightly in apical half, apex slightly to strongly asymmetrical and pointing toward midline of tegmen; in lateral view length/width ratio 3.4–4.6, setae confined to apex with length Female. Similar to male, no sexual dimorphism observed. Length 2.59–3.51 mm, width 1.03–1.31 mm (n = 10). Head width 0.62–0.69 mm and 0.69–0.76× pronotal width; dorsal interocular distance 0.36–0.43 mm, head width/dorsal interocular distance ratio 1.60–1.67, ventral interocular distance 0.27–0.32 mm, head width/ventral interocular distance ratio 2.00–2.25. Pronotum length/width ratio 0.68–0.72. Elytra length/width ratio1.69–1.78. Genitalia (n=4): gonostylus cylindrical, widest at apex, tapering slightly and gradually to base, set apically on gonocoxite, length= 3.9–5.6× width (Fig. 6K). Spermatheca distinct from all other Pharaxonotha studied so far from throughout its range (Tang, unpublished data): compact and arcuate, but not C-shaped, length/width = 2.8–3.3, apical third approximately 1.3× width of basal third, apical half with moderate annulation, apex rounded, basal third with longitudinal striations, spermathecal and glandular duct adjacent and inserted in a shallow pit on dorsal side of the basal third (Fig. 6J). Type locality. Mexico, Chiapas, near the border of Reserva de Biosfera Selva el Ocote [GPS and locality name omitted], forest understory on limestone. Range. The original description of the host of this beetle species, Ceratozamia santillanii, indicates its range as the northern highlands of Chiapas State, including the Ocote Biosphere Reserve (Pérez-Farrera et al. 2009). Material examined. Holotype (by designation) female of Pharaxonotha perezi with the following labels: 1) [rectangular; white; printed in black ink] “ MEX., Chis, [GPS coord. and locality omitted], cone Ceratozamia santillanii ♂, 16-V-2021, H. Gomez Dominguez & M. Jones ”; 2) [rectangular; red; printed in black ink] HOLOTYPE ♀ Pharaxonotha perezi Tang & Gomez Dominguez 2022. Deposited in the FSCA. Allotype (FSCA) and 14 adult paratypes ( same data as holotype) deposited at FSCA, IEXA. Additional specimens studied, but not included in types series: Mexico: Hidalgo, Hwy 105, La Mora, [GPS omitted], 1289m, Ceratozamia chamberlainii mid shed ♂ cone, 23-IV-2018, M. A. Perez Farrera & W. Tang (10); San Luis Potosí, Hwy 70, E of Tamasopo, 640m, [GPS omitted], cone Ceratozamia microstrobila [C. latifolia] ♂, 21-IV-2018, M. A. Perez Farrera & W. Tang (57); Potrero de Carneros Km 17 Carr. Rio Verde-Cd. Valles [GPS omitted], 900m, J. A. Peréz de la Rosa y G. Vargas cols. Ex Ceratozamia latifolia (estrobio masculine), 4.VII.2003 (4); Tampermoche, [GPS omitted], ex ♂ Ceratozamia hildae late shed cone, 19-IV-2018, M. A. Perez Farrera & W. Tang (20). These will be deposited in the institutions stated above. Etymology. Named in honor of Miguel Angel Pérez-Farrera for his work on the systematics, ecology and conservation of Mexican cycads. Remarks. A female specimen was chosen as the holotype due to the distinctive and highly diagnostic spermatheca of the female in this species. Current morphological analysis did not reveal differences in size or body proportions between the P. perezi population found on C. santillanii in Chiapas and similar beetles collected from C. chamberlainii, C. hildae and C. latifolia in Hidalgo and San Luis Potosi states, over 500 km to the northwest. Therefore, it is unclear whether beetles from these two regions represent disjunct isolated species or whether they represent the two ends of a more continuous distribution across this range. Dwarf Ceratozamia species, similar to these four, occur on similar steep habitats in Veracruz state (Vovides et al. 2008) between these two areas, presenting the possibility of a more continuous distribution for this beetle. Survey of dwarf Ceratozamia species in Veracruz for cone beetles and analysis of DNA may help to resolve the identity of these similar northern Pharaxontha beetles. Here the type series for P. perezi is restricted to C. santillanii., Published as part of Tang, William & Dominguez, Hector Gomez, 2022, Two new species of Pharaxonothinae beetles (Coleoptera: Erotylidae) inhabiting cones of the cycad Ceratozamia santillanii Pérez-Farr. & Vovides (Cycadales: Zamiaceae) in Mexico, pp. 428-442 in Zootaxa 5150 (3) on pages 437-439, DOI: 10.11646/zootaxa.5150.3.7, http://zenodo.org/record/6623165, {"references":["Tang, W., Xu, G., Marler, T., Khuraijam, J. S., Singh, R., Lindstrom, A. J., Radha, P., Rich, S., Nguyen, K. S. & Skelley, P. (2020) Beetles (Coleoptera) in cones of cycads (Cycadales) of the northern hemisphere: diversity and evolution. Insecta Mundi, 0781, 1 - 19.","Perez-Farrera, M. A., Vovides, A. P., Martinez-Camilo, R., Martinez-Melendez, N. & Iglesias, C. (2009) A reassessment of the Ceratozamia miqueliana species complex (Zamiaceae) of southeastern Mexico, with comments on species relationships. Systematics and Biodiversity, 7, 433 - 443. https: // doi. org / 10.1017 / S 1477200009990211","Vovides, A. P., Avendano, S., Perez-Farrera, M. A. & Gonzalez-Astorga, J. (2008) A new species of Ceratozamia (Cycadales: Zamiaceae) from Veracruz, Mexico. Novon, 18, 109 - 114. https: // doi. org / 10.3417 / 2006063"]}

Pharaxonotha gigantea Skelley, Tang and Pérez-Farrera, new species Figures 6A–H Diagnosis. This is the largest known Pharaxonotha with a length of 5.08 mm. Other distinguishing characters include the wide, transversely rectangular pronotum (length/width = 0.71) with punctures large, impressed, sharply defined. Description. Length 5.08 mm, width 1.95 mm. General body color (Fig. 6A–C) red-brown, pronotum slightly darker than elytra; dorsal surface punctate, with a procumbent hair associated with each punctation on head, pronotum and elytra, length of hairs reaching or nearly reaching nearest puncture (many hairs may have been broken off during the cleaning of the specimen, which was originally encased in pollen and other debris). Head. Not broad, width = 0.68× pronotal width (Fig. 6D–E); in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, finely, densely punctured, average distance between closest punctures 1–2× width of puncture; head width 1.07 mm; dorsal interocular distance 0.68 mm, head width/dorsal interocular distance ratio 1.57, ventral interocular distance 0.50 mm, head width/ventral interocular distance ratio 2.14. Eye with large black facets, about 2× diameter of head punctures. Antennal length slightly shorter than pronotal width, 1.2× head width; antennomere I (scape) fairly large, slightly elongate; antennomere II shorter than III; IV small, circular; V–VII same length as IV, gradually becoming wider with VIII transverse and flattened apically; club fairly large, IX and X similar in length; XI not enlarged, 1.3× longer than X, globular with rounded apex. Clypeus weakly concave anteriorly, moderately punctate. Transverse occipital line [vertexal line] distinct, laterally nearly reaching eye. Mentum and submentum finely punctured, ½–1/3× diameter of facet, distance between nearest punctures approximately 1× own diameter, each puncture with a short seta. Gular area smooth, without punctation or setae, border with submentum marked by change in punctation. Thorax. With pronotum transversely rectangular in dorsal view, length/width ratio 0.71; with distinct marginal carina laterally and basally, anteriorly with fine marginal carina medially; dorsally flattened; anterior angles narrowly rounded, projecting forward; posterior angles weakly developed, with small denticle at angle; lateral margin parallel-sided in medial half, shallowly arcuate inward anteriorly and posteriorly; posterior margin slightly projecting medially, projection beginning approximately by pair of small, dark pores in margin located ¼ width from posterior angles, each pore marks base of a distinct sulcus extending anteriorly onto disc ¼ length of pronotum. Prosternum in ventral view convex, with fine dense punctures; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae, longest setae approximately 1/3 length of eye; prosternal process flattened apically, weakly expanded and truncate at apex. Hypomeron rugose with dense moderate punctures. Scutellar shield distinctly transverse, posterior margin weakly roundly pentagonal. Elytra in dorsal view elongate, flattened dorsally; length/width 1.71, greatest width near midlength; with distinct marginal line basally; 10 complete striae of moderate puncture size; scutellary striole extending ¼ elytral length, with 15 punctures; punctures of elytral striae same size as pronotal punctures, striae weakly impressed; intervals of striae with fine, shallow punctures, ¼ size of strial punctures; all punctures of elytra bearing a single short seta; seta most visible in profile, extending to reach next puncture. Mesoventrite with strong coarse punctation, narrowly separated, hidden in rugose surface. Metaventrite glossy, with strong lateral punctation separated by 1–2× own diameter; medial surface finely distinctly punctured, separated by 3–4× own diameter; surface medially flattened, metathoracic discrimen extending approximately ¾ metaventrite length. Legs broadened, relatively similar in length and shape. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora robust, compressed laterally; tibiae shorter than femora, narrowly triangularly dilated to obliquely truncate apices; protibia with apical lateral tooth small, with complete apical fringe of very short stout spinules on straight ventral apical margin; meso- and metatibia with apical fringe of short stout spinules on anterior margin, finer setae on posterior margins. Abdomen. Ventrite apical margin bearing short, sparse setae; all ventrites bearing moderate, shallow punctation evenly distributed across surface, distance to nearest puncture approximately 2× diameter of puncture, punctures bearing mostly reclining setae; ventrite V with setae length nearly uniformly approximately 2× diameter of puncture; I–IV each with 2 or more median pairs of longer, semi-erect sensory hairs (difficult to see in poor lighting, often abraded). Male genitalia (n = 1): moderately dorsoventrally flattened; tegmen in dorsal view with basal piece ring-like and wide; parameres in dorsal view narrowing slightly 1/3 of length from the base then expanding gradually to a rounded apex, apical 2/3 slightly asymmetric with inner margin straight and outer margin with slight curvature, in lateral view length = 4.9× width; elongate cylindrical median lobe with strong taper apically; long coiled flagellum (Fig. 6F–H). Female. Not known. Distribution. Currently known only in the Mexican state of Guerrero on Dioon stevensonii. Material examined. Holotype (by designation) male of Pharaxonotha gigantea with the following labels: 1) [rectangular; white; printed in black ink] “ MEXICO, Guerrero, Achotla, oak forest, [GPS omitted], ex: Dioon tomasellii [D. stevensonii] male cone, coll. J. Chemnick, 18May2004, TW2004-18, elev. 1380 m; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Pharaxonotha gigantea Skelley, Tang and Pérez-Farrera 2022 ”. Deposited in the FSCA. Etymology. Named for its size; this is the largest Pharaxonotha known. Remarks. The sole type specimen of Pharaxonotha gigantea was found together on a male cone of D. stevensonii with a species of Allocorynina weevil, Parallocorynus (Dysicorynus) andrewsi Tang and O’Brien (O’Brien and Tang 2015). Two specimens of Pharaxonotha gigantea were originally collected, however, one was sacrificed for genetic analysis, but did not yield usable DNA., Published as part of Skelley, Paul E., Tang, William & Pérez-Farrera, Miguel Angel, 2022, Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting the cycad genus Dioon Lindl. (Cycadales), with descriptions of nine new species and comments on P. kirschii Reitter, pp. 1-41 in Insecta Mundi 2022 (917) on pages 17-19, DOI: 10.5281/zenodo.6391973, {"references":["O'Brien C, Tang W. 2015. Revision of the New World cycad weevils of the subtribe Allocorynina, with description of two new genera and three new subgenera (Coleoptera: Belidae: Oxycoryninae). Zootaxa 3970: 1 - 87."]}

Pharaxonotha Reitter Pharaxonotha Reitter 1875: 44. Type species. Pharaxonotha kirschii Reitter 1875: 44, by monotypy. Pakaluk 1988: 447 (review); Skelley and Tang 2020: 3 (key). Planismus Casey 1890: 500. Type species: Planismus floridanus Casey 1890: 500, by monotypy. Kingsolver 1973: 247 (synonymy); Pakaluk 1988: 447 (review); Skelley and Tang 2020: 3 (key). Diagnosis. Pharaxonotha can be distinguished from other members of the Pharaxonothinae by the following combination of characters: antennal club of 3 antennomeres; eyes large, encroaching upon gular area; submentalgular suture apparently lacking; stridulatory files at base of head separated by distance approximately equal to width of scutellar shield; anterior pronotal margin lacking or with very fine marginal line; lateral pronotal carina narrow, same thickness along entire length; elytra with basal bead; internal abdominal calli absent; male genitalia straight, not twisted, median lobe and tegmen rotated 180 degrees when retracted into the abdomen [see Fig. 10G and Tang et al. 2020 for examples; while it is common in the Cucujoidea for the male genitalia to be rotated 90 degrees while retracted into the abdomen (McHugh et al. 1997), 180 degrees is unusual]; median lobe cylindrical; flagellum long hair-like; tegmen flattened or not flattened, basal piece broadened and ring-like to hold coiled flagellum. Known distribution includes the Caribbean Basin from Florida and Bahamas to Greater Antillean islands of the Cayman Islands, Cuba, Jamaica, Hispaniola and Puerto Rico; mainland from southern US and Mexico to South America as far south as Bolivia., Published as part of Skelley, Paul E., Tang, William & Pérez-Farrera, Miguel Angel, 2022, Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting the cycad genus Dioon Lindl. (Cycadales), with descriptions of nine new species and comments on P. kirschii Reitter, pp. 1-41 in Insecta Mundi 2022 (917) on page 4, DOI: 10.5281/zenodo.6391973, {"references":["Reitter E. 1875. Revision der europaischen Cryptophagiden. Deutsche Entomologische Zeitschrift 19 (3): 1 - 86.","Pakaluk J. 1988. Review of the New World species of Pharaxonotha Reitter (Coleoptera: Languriidae). Revista de Biologia Tropical 36 (2 B): 447 - 451.","Skelley PE, Tang W. 2020. Two new species of Pharaxonotha Reitter among the early-diverging lineages, with a key to the species of the genus (Coleoptera: Erotylidae: Pharaxonothinae). Insecta Mundi 0837: 1 - 11.","Casey. T. L. 1890. Coleopterological notices II. Annals of the New York Academy of Sciences 5: 307 - 504.","Kingsolver JM. 1973. New synonymy in Languriidae (Coleoptera). Proceedings of the Entomological Society of Washington 75 (2): 247.","Tang W, Xu G Marler T, Khuraijam JS, Singh R, Lindstrom AJ, Radha P, Rich S, Nguyen KS, Skelley P. 2020. Beetles (Coleoptera) in cones of cycads (Cycadales) of the northern hemisphere: diversity and evolution. Insecta Mundi 0781: 1 - 19.","McHugh JV, Marshall CJ, Fawcett FL. 1997. A study of the adult morphology in Megalodacne heros (Say) (Coleoptera: Erotylidae). Transactions of the American Entomological Society 123: 167 - 223."]}

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Two new species of Pharaxonotha Reitter among the early-diverging lineages, with a key to the species of the genus (Coleoptera: Erotylidae: Pharaxonothinae). Insecta Mundi 0837: 1-11.","Skelley P, Xu G, Tang W, Lindstrom A, Marler T, Khuraijam JS, Singh R, Radha P, Rich S. 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: 1-63.","Tang W. 1987. Insect pollination in the cycad Zamia pumila (Zamiaceae). American Journal of Botany 74: 90-99.","Tang W, Skelley P, Perez-Farrera MA. 2018a. Ceratophila, a new genus of erotylid beetles (Erotylidae: Pharaxonothinae) inhabiting male cones of the cycad Ceratozamia (Cycadales: Zamiaceae). Zootaxa 4508: 151-178.","Tang W, Xu G, O'Brien CW, Calonje M, Franz NM, Johnston MA, Taylor A, Vovides AP, Perez-Farrera MA, Salas- Morales SH, Lazcano-Lara JC, Skelley P, Lopez-Gallego C, Lindstrom A, Rich S. 2018b. Molecular and morphological phylogenetic analyses of New World cycad beetles: what they reveal about cycad evolution in the New World. Diversity 10(38): 1-26.","Tang W, Xu G Marler T, Khuraijam JS, Singh R, Lindstrom AJ, Radha P, Rich S, Nguyen KS, Skelley P. 2020. Beetles (Coleoptera) in cones of cycads (Cycadales) of the northern hemisphere: diversity and evolution. Insecta Mundi 0781: 1-19.","Valencia-Montoya WA, Tuberquia D, Guzman PA, Cardona-Duque J. 2017. Pollination of the cycad Zamia incognita A. Lindstr. & Idarraga by Pharaxonotha beetles in the Magdalena Medio Valley, Colombia: a mutualism dependent on a specific pollinator and its significance for conservation. Arthropod-Plant Interaction 5: 717-729.","Vovides AP. 1991. 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Pharaxonotha thomasi Skelley and Tang, new species Figure 2A–I Diagnosis. Pharaxonotha thomasi is one of the most recognizable species of the genus. It is the only species where all tibiae are strongly triangularly dilated apically; protibia with a complete row of short stout spinules on a straight ventral apical margin; body distinctly flattened; and with a known distribution in Honduras on Zamia onan-reyesii. Description. Length 2.44–2.96 mm, width 0.96–1.19 mm. Body (Fig. 2A–C) in dorsal view elongate, distinctly flattened, greatest width at middle of elytra; in lateral view flattened dorsally. General body color entirely red-brown; dorsal surface punctate, shining and appearing glabrous, short procumbent hairs associated with punctation on pronotum and elytra, ventrally shining and appearing glabrous except mesoventrite and abdomen mostly covered with short procumbent setae. Head. Not broad, width = 0.68–0.74× pronotal width; in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, finely, moderately punctured, average distance between closest punctures 2–3× width of puncture; head width 0.56–0.65 mm; dorsal interocular distance 0.33–0.37 mm, head width/dorsal interocular distance ratio 1.63–1.80, ventral interocular distance 0.21–0.26 mm, head width/ventral interocular distance ratio 2.55–2.89. Eye with large black facets, about 2× diameter of head punctures. Antennal length slightly shorter than pronotal width, 1.2× head width; antennomere I (scape) fairly large, slightly elongate; antennomere II equal in size to III; IV small, circular; V–VII same length as IV, gradually becoming wider with VIII distinctly transverse and flattened apically; club fairly large, IX and X similar in length; XI not enlarged, 1.4× longer than X, globular with rounded apex. Clypeus weakly concave anteriorly, moderately punctate. Transverse occipital line [vertexal line] distinct from eye to eye. Mentum and submentum somewhat coarsely punctured, 2–3× diameter of facet, distance between nearest punctures approximately 1× own diameter, each puncture with a short seta. Gular area smooth, without punctation or setae, border with submentum marked by change in punctuation. Thorax. With pronotum transversely rectangular in dorsal view, length/width ratio 0.67–0.73; with distinct marginal carina laterally and basally, anteriorly with fine marginal carina medially; dorsally flattened; anterior angles broadly rounded, not projecting forward; posterior angles weakly developed, with small denticle at angle; lateral margin parallel-sided in medial half, shallowly arcuate inward anteriorly and posteriorly; posterior margin slightly projecting medially, projection beginning approximately by pair of small, dark pores in margin located 1/4 width from posterior angles, each pore marks base of a distinct sulcus extending anteriorly onto disc 1/4 length of pronotum. Prosternum in ventral view convex, with few scattered punctures; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae, longest setae approximately 1/3 length of eye; prosternal process flattened apically, expanded and truncate at apex. Hypomeron laterally with few minute punctures, medially lacking distinct longitudinal striations. Scutellar shield distinctly transverse, posterior margin weakly roundly pentagonal. Elytra in dorsal view elongate, flattened dorsally; length/width 1.44–1.62, greatest width near midlength; with distinct marginal line basally; 10 complete striae of moderate puncture size; scutellary striole extending 1/4 elytral length, with 10–15 punctures; punctures of elytral striae 1.5× larger than pronotal punctures, striae weakly impressed; intervals of striae with fine, shallow punctures, 1/4 size of strial punctures; all punctures of elytral bearing a single short seta; seta only visible in profile, extending slightly out of puncture. Mesoventrite with strong punctation, distance between nearest punctures approximately equal to diameter of punctures, puncture depth moderate. Metaventrite glossy, with strong lateral punctation separated by 1–2× own diameter; medial surface finely distinctly punctured, separated by 3–4× own diameter; surface medially flattened, metathoracic discrimen extending approximately 3/4 metaventrite length. Legs broadened, relatively similar in length and shape. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora robust, compressed laterally; tibiae shorter than femora, triangularly dilated to obliquely truncate apices, obliquely truncate apical margin width 1/3 tibial length; protibia with apical lateral tooth small, with complete apical fringe of very short stout spinules on straight ventral apical margin; meso- and metatibia with apical fringe of short stout spinules on anterior margin, finer setae on posterior margins. Abdomen. Ventrite apical margin bearing short, sparse setae; all ventrites bearing moderate, shallow punctation evenly distributed across surface, distance to nearest puncture approximately 2× diameter of puncture, punctures bearing mostly reclining setae; ventrite V with setae length nearly uniformly approximately 2× diameter of puncture; I–IV each with 2 or more median pairs of longer, semi-erect sensory hairs (difficult to see in poor lighting, often abraded). Male genitalia (n = 2) similar to all others in the genus, with dorsoventrally flattened tegmen, elongate cylindrical median lobe, and long coiled flagellum (Fig. 2D–G). Female. Similar to male, sexual dimorphism weakly evident with male protarsomere I more broadly dilated. Genital tube shortened, length past abdominal segment VIII = 1.5× width (n = 2); gonostylus set apically on gonocoxite, gonostylus length = 4–5× width (Fig. 2H). Spermatheca shaped like a pill capsule, length approximately 3× width, basal half smooth, apical half with reticulation, folding at center (Fig. 2I). Range. Known from Honduras, the type locality, on male cones of Zamia onan-reyesii. Material examined. Holotype (by designation) male of Pharaxonotha thomasi with the following labels: 1) [rectangular; white; printed in black ink] “ HONDURAS: San Pedro Sula, Filo del Cerrito, Laguna Tembladeros, 30-VII-2003, [GPS coord. omitted], col. Jody Haynes, on male cone of Zamia sp. ” 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Pharaxonotha thomasi Skelley & Tang 2020 ”. Deposited in the FSCA. Female allotype and adult paratypes (n = 41): same data as holotype (4 ANIC; allotype, 25 FSCA; 4 NHMUK; 4 NZAC; 4 USNM). In Tang et al. (2020), this species is presented on their tree as “D0066 Z. onanreyesii > HONDURAS ”. Etymology. In the early 1990s, Andrew Vovides sent specimens from a Ceratozamia cone to Michael C. Thomas for identification, who recognized them as multiple undescribed taxa. While Mike was an expert on cucujoid beetles, he had not published on this group of beetles. James Pakaluk, who had recently described species in the group (Pakaluk 1988), agreed to describe them. However, Pakaluk left entomological work in the late 1990s and the task of describing the new taxa fell back to Thomas. In the early 2000s, Węgrzynowicz (2002) and Leschen (2003) independently merged several higher taxa into the family Erotylidae, including Pharaxonotha. At that time, author PES agreed to help Mike Thomas with the descriptions. With this new enthusiasm additional materials began arriving from cycad researchers, most representing obviously undescribed taxa or presenting taxonomic problems that needed more detailed study before being described. Mike eventually turned the entire project over to PES, who was quickly overwhelmed with the diversity and complexity, but was able to help others describe individual species (Chaves and Genaro 2005; Franz and Skelley 2008). Around 2010, Willie Tang, Guang Xu, and others began work on relationships of cycad pollinating beetles using molecular methods. A team formed and progress on “ Pharaxonotha ” finally started (Skelley 2013; Xu et al. 2015; Skelley et al. 2017; Tang et al. 2018a, 2018b, 2020; Skelley and Segalla 2019). After his initial 1990s work, Mike Thomas encouraged and helped others working on this complex genus, until he died in October 2019. It is an honor that we recognize Mike’s enthusiasm and encouragement of others to study beetles by naming this species after him. If not for his initial push, a lot of this work would not have happened. Remarks. The host for P. thomasi is Zamia onan-reyesii (Nelson and Sandoval 2008, Schutzman et al. 2008), an arborescent species of the Mesoamerica clade of Zamia as identified via genetic analysis by Calonje et al. (2019)., Published as part of Skelley, Paul E. & Tang, William, 2020, Two new species of Pharaxonotha Reitter among the early-diverging lineages, with a key to the species of the genus (Coleoptera: Erotylidae: Pharaxonothinae), pp. 1-11 in Insecta Mundi 2020 (837) on pages 7-9, DOI: 10.5281/zenodo.5353903, {"references":["Tang W, Xu W, Marler T, Khuraijam JS, Singh R, Lindstrom AJ, Radha P, Rich S, Nguyen KS, Skelley P. 2020. Beetles (Coleoptera) in cones of cycads (Cycadales) of the northern hemisphere: diversity and evolution. Insecta Mundi 0781: 1 - 19.","Pakaluk J. 1988. Review of the New World species of Pharaxonotha Reitter (Coleoptera: Languriidae). Revista de Biologia Tropical 36 (2 B): 447 - 451.","Wegrzynowicz P. 2002. Morphology, phylogeny and classification of the family Erotylidae based on adult characters (Coleoptera: Cucujoidea). Genus 13 (4): 435 - 504.","Leschen RAB. 2003. Erotylidae (Insecta: Coleoptera: Cucujoidea): phylogeny and review. Fauna of New Zealand No. 47. Manaaki Whenua Press; Lincoln, NZ. 103 p.","Chaves R, Genaro JA. 2005. A new species of Pharaxonotha (Coleoptera: Erotylidae), probable pollinator of the endangered Cuban cycad, Microcycas calocoma (Zamiaceae). Insecta Mundi 19 (3): 143 - 150.","Franz NM, Skelley PE. 2008. Pharaxonotha portophylla (Coleoptera: Erotylidae), new species and pollinator of Zamia (Zamiaceae) in Puerto Rico. Caribbean Journal of Science 44 (3): 321 - 333.","Skelley P. 2013. Henoticonus bouchardi Grouvelle transferred to Trogocryptoides Champion (Coleoptera: Salpingidae). Insecta Mundi 0281: 1 - 4.","Xu G, Tang W, Skelley P, Liu N, Rich S. 2015. Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa 3986: 251 - 278.","Skelley P, Xu G, Tang W, Lindstrom A, Marler T, Khuraijam JS, Singh R, Radha P, and Rich S. 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 - 63.","Tang W, Skelley P, Perez-Farrera MA. 2018 a. Ceratophila, a new genus of erotylid beetles (Erotylidae: Pharaxonothinae) inhabiting male cones of the cycad Ceratozamia (Cycadales: Zamiaceae). Zootaxa 4508: 151 - 178.","Tang W, Xu G, O'Brien CW, Calonje M, Franz NM, Johnston MA, Taylor A, Vovides AP, Perez-Farrera MA, Salas- Morales SH, Lazcano-Lara JC, Skelley P, Lopez-Gallego C, Lindstrom A, Rich S. 2018 b. Molecular and morphological phylogenetic analyses of New World cycad beetles: what they reveal about cycad evolution in the New World. Diversity 10 (38): 1 - 26.","Skelley P, Segalla R. 2019. A new species of Pharaxonotha Reitter (Coleoptera: Erotylidae) from central South America. Zootaxa 4590 (1): 184 - 190.","Nelson C, Sandoval Gonzales GG. 2008. Una especie nueva de Zamia (Zamiaceae) de Honduras. Ceiba 49: 135 - 136.","Schutzman B, Adams R, Haynes JL, Whitelock LM. 2008. A new endemic Zamia from Honduras (Cycadales: Zamiaceae). The Cycad Newsletter 31 (2 / 3): 22 - 25.","Calonje M, Meerow AW, Griffith MP, Salas-Leiva D, Vovides AP, Coiro M, Francisco-Ortega J. 2019. A time-calibrated species tree phylogeny of the New World cycad genus Zamia L. (Zamiaceae, Cycadales). International Journal of Plant Sciences 180: 286 - 314."]}

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Key to genera of Pharaxonothinae (modified from Leschen 2003 and Xu et al. 2015) 1 Antennal club of 2 antennomeres.......................................................... Setariola Jacobson - Antennal club of 3 antennomeres......................................................................... 2 2 Pronotal callosities present; vestiture of scale-like setae................................ Leucohimatium Rosenhauer - Pronotal callosities absent; vestiture of simple setae.......................................................... 3 3 Eyes small, ventrally not encroaching upon gular area, distance between eye and lateral corner of mentum about equal to length of eye in ventral view; not associated with cycads...................................................... 4 - Eyes large, ventrally encroaching upon gular area, distance between eye and lateral corner of mentum 5 4 Dorsal punctation strong and impressed.................................................... Henoticonus Reitter - Dorsal punctation weak and not strongly impressed......................................... Loberogosmus Reitter 5 Submental-gular suture visible as depressed darkened area, often with row of punctures and setae; anterior pronotal margin of most with fine, distinct, complete marginal line; male genitalia with median lobe and tegmen laterally flattened and slightly to strongly twisted (at least basally); spiculum gastrale asymmetrical; Asia............... Cycadophila Xu, Tang & Skelley - Submental-gular suture apparently lacking, surface flattened; anterior pronotal margin lacking marginal line, some with row of fine punctures; male genitalia with median lobe cylindrical and tegmen dorsoventrally or laterally flattened, but usually not twisted; spiculum gastrale symmetrical; Americas............................................................ 6 6 Lateral pronotal carina thicker anteriorly in lateral view (Figs. 1 D���E); head with supraocular lines and elytra lacking basal bead; stridulatory file at base of head separated by distance ~ 0.5�� width of the scutellar shield (Fig. 1B); tarsomeres I���III distinctly widened (Figs. 4 D���E, 8F), metatarsomere I width �� length; male genitalia with median lobe arched in lateral view; flagellum length Šlength of median lobe and not coiled (Figs. 4G, 5F, 6H, 7G, 8G); Mexico and probably Belize, Guatemala and Honduras on Ceratozamia............................................................ Ceratophila, new genus - Lateral pronotal carina narrow, same thickness along entire length in lateral view (Fig. 1C); head without supraocular lines and elytra with distinct basal bead (Fig. 1A; P. esperanzae Chaves and Genaro is the exception); stridulatory files at base of head separated by distance ~ width of scutellar shield (Fig. 1A); tarsomeres I���III narrow, metatarsomere I length always distinctly greater than width; male genitalia with median lobe straight; flagellum length >> length of median lobe and coiled; widespread New World on Ceratozamia, Dioon, Microcycas, Zamia..................................... Pharaxonotha Reitter, Published as part of Tang, William, Skelley, Paul & P��rez-Farrera, Miguel Angel, 2018, Ceratophila, a new genus of erotylid beetles (Erotylidae: Pharaxonothinae) inhabiting male cones of the cycad Ceratozamia (Cycadales: Zamiaceae), pp. 151-178 in Zootaxa 4508 (2) on page 153, DOI: 10.11646/zootaxa.4508.2.1, http://zenodo.org/record/2606934, {"references":["Leschen, R. A. B. (2003) Fauna of New Zealand 47. Erotylidae (Insecta: Coleoptera: Cucujoidea): Phylogeny and Review. Manaaki Whenua Press, Lincoln, 108 pp.","Xu, G., Tang, W., Skelley, P., Liu, N. & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278. https: // doi. org / 10.11646 / zootaxa. 3986.3.1"]}

Cycadophila (Strobilophila) 16(1). Protibia notably dilated toward apex with stout marginal spines on lateral apical 1/3 separated by less than their length (Figs. 2 F, 20D); with distinct furrow on each side of pronotal base (Fig. 19 A)........................................ 17 - Protibia weakly dilated distally with stout marginal spines on lateral apical 1/2 separated by their length or more (Figs. 2 G, 21D); 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, 19G); 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, 19A); 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, Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 11, DOI: 10.5281/zenodo.575641

Cycadophila (Strobilophila) assamensis Skelley, Xu & Tang, new species Figs. 16 A���G 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 16S 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. 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 �� 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. 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, NG1���; 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. Etymology. Named for the State where the type specimens were collected. 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 t1 tubercle (see Key to larvae above) vs. the vertically oriented t1 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 48-50, DOI: 10.5281/zenodo.575641, {"references":["Singh, K. J. & Singh, R. (2014) Population assessment and distribution of Cycas pectinata"]}

Cycadophila (Cycadophila) papua Skelley, Xu & Tang, new species Figs. 13 A–E 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. 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. 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. Etymology. Named for the region of New Guinea the types are from, Papua is used as a noun in apposition. 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 1994a) and another species, C. rumphii, occurs along coastal habitats at lower elevations (Hill 1994b).

Key to adults of Cycadophila 1. Lateral pronotal carinae narrow in lateral view, with carinal punctures in a single row (Figs. 2 A,C); protibia dilated or not distally, lacking stout setae along lateral margin of protibia or with few setae at extreme apex (Figs. 2 D���E) (may have stout setae along ventro-apical margin) [Subgenus Cycadophila Xu, Tang & Skelley]........................................ 2 - Lateral pronotal carinae thickened in lateral view, with carinal punctures scattered (Fig. 2 B); protibia dilated distally with 5���6 short stout setae along apical third of lateral margin (Figs. 2 F���G) [Subgenus Strobilophila, new subgenus]............ 16 2(1). Head dorsally flattened, clypeus flat in lateral view (Fig. 2 A); ventral interocular distance 0.22-0.45�� head width; prosternal process narrow (1/2 width of profemur with apex truncate; pronotal angles sharply angulate, surface near angle reflexed, lateral carinae parallel most of length, basally appearing quadrate with basal margin angled where discal groove meets margin; unless teneral, body dark brown ventrally, elytra usually without distinct color pattern [nigra species group] ��� 8 3(2). Protibia narrow distally, with apical fringe of fine distally directed setae, without stout spinules (Fig. 2 D); male genitalia moderate in length, weakly twisted.......................................................................... 4 - Protibia dilated distally (distinctly in males*, less so in females), with apical fringe of short, stout, ventrally directed spinules (Fig. 2 E); male genitalia short and distinctly twisted [lata species group]........................................ 6 [*Note: Tibial and femoral dilation is sexually dimorphic in the lata group, and there is variation in males from major to minor leg development. Minor males or females may not key well beyond this point. In general, male protibiae are distinctly dilated in comparison to metatibiae, while female pro- and metatibiae are similarly dilated.] 4(3). Supraocular stria long, nearly full length of eye; terminal maxillary palpomere longer than entire length of labial palp; pronotum black, contrasting with orange-brown venter [papua species group].......................................... 5 - Supraocular striae short, less than half length of eye; terminal maxillary palpomere shorter than entire length of labial palp; pronotum same orange-brown color as venter [fupingensis species group]..................................................................................................... Cycadophila (C.) fupingensis Skelley, Tang & Xu 5(4). Elytral orange marks broader, broadly reaching base between striae III���VIII; prosternal process with coxal lines parallel; Philippines....................................................................... C. (C.) samara, new species - Elytral orange marks narrower, narrowly reaching base between striae V���VI; prosternal process with coxal lines diverging posteriorly; Papua New Guinea...................................................... C. (C.) papua, new species 6(3). Head width/ventral interocular distance ratio>4.00, eyes narrowly separated ventrally; width of elytra greatest anterior to middle; male mesotibia strongly dilated, lateral margin arched (Fig. 2 J), female unknown; ���Tonkin���.... C. (C.) lata (Grouvelle) - Head width/ventral interocular distance ratio Cycas debaoensis in Guangxi, China & C. hoabinhensis in northern Vietnam............................................................................... C. (C.) debaonica Xu, Tang & Skelley - Most lacking (or at most with an indistinct, obtuse) fine lateral projection behind pore at posterior pronotal angle (Fig. 2 K); major male mesofemora with posterior margin straight medially, abruptly turning to another flat margin apically (Fig. 2 O); inhabiting Cycas collina & C. tanqingii in Yunnan, China, northern Laos and northern Vietnam...................................................................................................... C. (C.) collina , new species, Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 10, DOI: 10.5281/zenodo.575641

Cycadophila (Strobilophila) kwaiensis Skelley, Xu & Tang, new species Figs. 18 A���E 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 16S rRNA mitochondrial gene (GenBank Accession number KY365221). 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. 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. 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. Etymology. Named for the locality of collection in the vicinity of the Kwai River Bridge in Kanchanaburi Province, Thailand. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 53-55, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Cycadophila (Strobilophila) yangi Skelley, Xu & Tang, new species Figs. 2 G, 21A–E 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. 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. 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, 640m, 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, 700m, 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. 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. 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 16S 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 D0333 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.

Cycadophila (Cycadophila) cyclochasma Skelley, Xu & Tang, new species Figs. 8 A���E 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. 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 90o; 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 45o 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. 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; 100m, 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. 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. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 27-29, DOI: 10.5281/zenodo.575641, {"references":["Osborne, R., Hill, K. D., Nguyen, H. T. & Ke, L. P. (2007) Cycads of Fietnam. Tien Wah Press (Pte) Ltd, Singapore, 116 pp."]}

Cycadophila (Strobilophila) Skelley, Xu & Tang, new subgenus Type species. Cycadophila (Strobilophila) tansachai, n. sp., here designated. 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���2.40mm. 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. Larva. Description based on a larval morphotype found with adults of Cycadophila (Strobilophila) tansachai in Cycas elephantipes male cones. 16S 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 T2���3 and A1���A7, anterior 3/4 of A8, central 1/2 of T1 and all of A9 pigmented dark brown, sclerotized; brown areas, except for anterior 1/4���1/3, more or less randomly covered with granules; sclerotized regions of T1���3 and A1���8 distinctly bisected by light-colored middorsal line; T2���3 and A1���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; A9 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 t3 (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. 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. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 45-48, DOI: 10.5281/zenodo.575641, {"references":["Tang, W., Oberprieler, R. & Yang, S. L. (1999) Beetles (Coleoptera) in Asian Cycas: diversity, evolutionary patterns, and implications for taxonomy. In: Chen, C. J. (Ed.), Biology and Conservation of Cycads. International Academic Publishers, Beijing, pp. 280 - 297.","Leschen, R. A. B. (2003) Erotylidae (Insecta: Coleoptera: Cucujoidea): phylogeny and review. Fauna of New Zealand, 47, 1 - 108.","Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Fupingensis species group 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 (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)., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 21, DOI: 10.5281/zenodo.575641

Nigra species group 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 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. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 22, DOI: 10.5281/zenodo.575641, {"references":["Hinton, H. E. (1945) A Monograph of the Beetles Associated with Stored Products. Folume 1. British Museum (Natural History), London, 443 pp."]}

Cycadophila (Cycadophila) lata (Grouvelle), new combination Figs. 2 J, 5A���F Pharaxonotha lata Grouvelle 1916: 61; Schenkling 1923: 15; Schenkling 1928: 31. 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. 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. 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). 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 1A, built by the French in the early 20th 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 19-21, DOI: 10.5281/zenodo.575641, {"references":["Grouvelle, A. (1916) Descriptions d'especes nouvelles de Cryptophagidae. Memoires Entomologiques. Etudes sur les Coleopteres, 1, 30 - 79.","Schenkling, S. (1923) Cryptophagidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 76, pp. 1 - 92.","Schenkling, S. (1928) Languriidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 100, 40 pp.","Osborne, R., Hill, K. D., Nguyen, H. T. & Ke, L. P. (2007) Cycads of Fietnam. Tien Wah Press (Pte) Ltd, Singapore, 116 pp.","Averyanov, L. V., Nguyen, T. H., Sinh, K. N., Pham, T. V., Lamxay, Bounphanmy, V. S., Lorphengsy, S., Phan, L. K., Lanorsavanh, S. & Chantthavongsa, K. (2014) Gymnosperms of Laos. Nordic Journal of Botany, 32, 765 - 805. https: // doi. org / 10.1111 / njb. 00498"]}

Cycadophila (Strobilophila) hiepi Skelley, Xu & Tang, new species Figs. 17 A���F 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. 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. 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., 140m, 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., 140m, 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, 66m, 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. 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. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 50-53, DOI: 10.5281/zenodo.575641

Papua species group 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 (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)., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 41, DOI: 10.5281/zenodo.575641, {"references":["Lindstrom, A. J., Hill, K. D. & Stanberg, L. C. (2009) The genus Cycas (Cycadaceae) in Indonesia. Telopea, 12 (3), 385 - 418. https: // doi. org / 10.7751 / telopea 20095826"]}

Cycadophila (Cycadophila) intermedia (Ch��j��), new combination (incertae sedis) Pharaxonotha intermedia Ch��j�� 1967: 39. Adult description. No specimen studied, Ch��j�����s (1967) description and key characters are repeated here: ���Length 5.0 0 mm. Body elongate-oval, lightly convex on dorsum. Head, together with the with basal segments of antennae, dark piceous, but the median area of frons more or less blackish, lustrous, and the antennal clubs light brownish; pronotum pitchy black, with the lateral areas broadly rather dark red-brown (this color broadened forwardly from the base), lustrous; scutellum shiny black; elytra black, lustrous, with the laterio-anterior and posterior areas rather dark red-brown; underside red-brown, with the median areas rather darker than the lateral areas; legs deep redbrown. 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; 1st segment strongly thickened and dilated terminally, 2nd segment as broad as it is long, 4th~7th segments moniliform, 3rd segment distinctly longer than wide, longer than either the 2nd and 4th segments, 8th segment transverse and broader than the 7th segment but far smaller than the 9th segment, 9th and 10th segments strongly enlarged with the former one semicircular and the latter transverse, 11th segment transverse subovate and distinctly wider than long, 11th segment narrower than either the 10th and 9th segments but very much larger than the 8th 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 1st tarsal segment not flattened.��� Type locality. Laos. Type. According to Ch��j�� (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.���. Ch��j�� (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. Remarks. Ch��j�����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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 34-35, DOI: 10.5281/zenodo.575641, {"references":["Chujo, M. (1967) Languriid-Beetles from Thailand, Laos, Viet-Nam, S. China, Is. Hainan, Formosa and Ryukyus. Studies on the Languriid-Beetles (12). Memoirs of the Faculty of Education Kagawa University, 2 (149), 1 - 42.","Skelley, P. E. (2013) Henoticonus bouchardi Grouvelle transferred to Trogocryptoides Champion (Coleoptera: Salpingidae). Insecta Mundi, 0 281, 1 - 4.","Averyanov, L. V., Nguyen, T. H., Sinh, K. N., Pham, T. V., Lamxay, Bounphanmy, V. S., Lorphengsy, S., Phan, L. K., Lanorsavanh, S. & Chantthavongsa, K. (2014) Gymnosperms of Laos. Nordic Journal of Botany, 32, 765 - 805. https: // doi. org / 10.1111 / njb. 00498"]}

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

Cycadophila (Cycadophila) yunnanensis (Grouvelle) Pharaxonotha yunnanensis Grouvelle 1916: 61; Schenkling 1923: 15; Mader 1926: 721; Schenkling 1928: 31; Węgrzynowicz 2007: 536. Cycadophila yunnanensis (Grouvelle); Xu et al. 2015: 20 ���23, 25���26 (lectotype designation). 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). 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. 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 16S 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)., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 40, DOI: 10.5281/zenodo.575641, {"references":["Grouvelle, A. (1916) Descriptions d'especes nouvelles de Cryptophagidae. Memoires Entomologiques. Etudes sur les Coleopteres, 1, 30 - 79.","Schenkling, S. (1923) Cryptophagidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 76, pp. 1 - 92.","Mader, L. (1926) Clavicornia, Cryptophagidae. In: Winkler, A. (Ed.), Catalogus Coleopterorum regionis palaearcticae, Pars 6, pp. 720 - 730.","Schenkling, S. (1928) Languriidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 100, 40 pp.","Wegrzynowicz, P. (2007) Family Erotylidae Latreille, 1802. In: Lobl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera, Folume 4. Apollo Books, Stenstrup, pp. 531 - 546.","Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Cycadophila (Cycadophila) samara Skelley, Xu & Tang, new species Figs. 14 A���E 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. 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. 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]. 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. Remarks. A specimen yielding a DNA sequence of the 16S rRNA gene that closely matches that of the type specimens was collected in Panay Island, Phillipines on Cycas edentata (see D0081 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 43-45, DOI: 10.5281/zenodo.575641

Cycadophila (Cycadophila) eurynota Skelley, Xu & Tang, new species Figs. 10 A���E 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. 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 90o; 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. 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. 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 ���. 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, Ch��j�� (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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 32-33, DOI: 10.5281/zenodo.575641, {"references":["Chujo, M. (1967) Languriid-Beetles from Thailand, Laos, Viet-Nam, S. China, Is. Hainan, Formosa and Ryukyus. Studies on the Languriid-Beetles (12). Memoirs of the Faculty of Education Kagawa University, 2 (149), 1 - 42.","Tang, W., Yang, S. and Vatcharakorn, P. (1997) Cycads of Thailand. Nong Nooch Tropical Garden, Bangkok, 34 pp."]}

Lata species group 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 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 15, DOI: 10.5281/zenodo.575641

Cycadophila (Cycadophila) nigra (Gorham) Thallis ? nigra Gorham 1895: 324. Pharaxonotha nigra (Gorham); Arrow 1925: 255; Schenkling 1928: 31; Ch��j�� 1964: 224, pl. 14, fig. 7; Ch��j�� 1967: 41. Pharaxonotha indica Grouvelle 1903: 127; Grouvelle 1916: 62 (sp. bona); Schenkling 1923: 15 (sp. bona); Arrow 1925: 255 [syn.]; Schenkling 1928: 31; Ch��j�� 1964: 224. Cycadophila nigra (Gorham); Xu et al. 2015: 17 ���20 (lectotype designation). 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). 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, 955m, 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���100m, 25-IV-2015, N. S. Khang, ex �� Cycas hoabinensis cone (NSK 795 b) (1). This additional material examined is deposited at ANIC and FSCA. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 35, DOI: 10.5281/zenodo.575641, {"references":["Gorham, H. S. (1895) List of the Coleoptera in the collection of H. E. Andrewes Esq. from India and Burma, with descriptions of new species and notes. Families: Malacodermata- Erotylidae- Endomychidae. Annales de la Societe Entomologique de Belgique, 39, 293 - 330.","Arrow, G. J. (1925) Coleoptera. Clavicornia. Erotylidae, Languriidae, and Endomychidae. In: Shipley, A. E. & Scott, H. (Eds.), The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, XVI + 416 pp., 76 figs., pl., map.","Schenkling, S. (1928) Languriidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 100, 40 pp.","Chujo, M. (1964) Coleoptera from Southeast Asia (III). 20. Family Languriidae. Nature and Life in Southeast Asia, 3, 219 - 226, pl. 14.","Chujo, M. (1967) Languriid-Beetles from Thailand, Laos, Viet-Nam, S. China, Is. Hainan, Formosa and Ryukyus. Studies on the Languriid-Beetles (12). Memoirs of the Faculty of Education Kagawa University, 2 (149), 1 - 42.","Grouvelle, A. (1903) Coleopteres de l'Inde recoltes par MM .. H. - L. Andrewes et T. - R. - D. Bell. Annales de la Societe Entomologique de France, 72, 125 - 127.","Grouvelle, A. (1916) Descriptions d'especes nouvelles de Cryptophagidae. Memoires Entomologiques. Etudes sur les Coleopteres, 1, 30 - 79.","Schenkling, S. (1923) Cryptophagidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 76, pp. 1 - 92.","Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Cycadophila (Cycadophila) Xu, Tang, Skelley C. (C.) lata species group 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 (Ch��j��) 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 16S 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 (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 (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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 8, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Cycadophila (Strobilophila) tansachai Skelley, Xu & Tang, new species Figs. 2 B, 2F, 2N, 19A���I, 20A���E 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 16S rRNA mitochondrial gene (GenBank Accession numbers KY356222 ��� KY365225, KY356244 ��� KY365245). 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. 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. 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, E101��27.260, 862m, 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, 640m, 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��53.835N, 101��27.260E, 862m, 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��38.732N, 101��23.483E, 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, 700m, 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��07.15N, 101��56.00E, 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, 370m, 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, 370m, 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. 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. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 55-58, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278.","Yang, S. - L. & Meerow, A. W. (1996) The Cycas pectinata (Cycadaceae) complex: genetic structure and gene flow. International Journal of Plant Science, 157, 468 - 483.","Lindstrom, A. J., Hill, K. D. (2002) New species and a new record of Cycas (Cycadaceae) in Thailand. Brittonia, 54, 298 - 304. https: // doi. org / 10.1663 / 0007 - 196 X (2003) 54 [298: NSAANR] 2.0. CO; 2","Singh, K. J. & Singh, R. (2014) Population assessment and distribution of Cycas pectinata"]}

Cycadophila (Cycadophila) fupingensis Skelley, Tang & Xu Figs. 2 A, 2D, 3E���H Cycadophila fupingensis Skelley, Tang & Xu in Xu et al. 2015: 12 ���17. 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). 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, 955m, 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���100m, 25-IV-2015, N. S. Khang ex ♂ Cycas hoabinhensis (1). These are deposited at ANIC and FSCA. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 21-22, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278.","Xie, J., Jian, S., & Liu, N. (2005) Genetic variation in the endemic plant Cycas debaoensis on the basis of the ISSR analysis. Australian Journal of Botany, 53, 141 - 145.","Fang, S. (2009) Study on Conservated Biology of Cycas debaoensis. Ph. D. dissertation, Sun Yat-Sen University, Guangzhou, 113 pp."]}

Cycadophila Xu, Tang & Skelley, 2015 Type species. Cycadophila debaonica Xu, Tang & Skelley, 2015, by original designation. 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. 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 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on page 9, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278."]}

Cycadophila (Cycadophila) discimaculata (Mader), new combination Figs. 9 A���F Pharaxonotha discimaculata Mader 1939: 46; Węgrzynowicz 2007: 536. Pharaxonata [sic] discimaculata: Mader 1955: 65. 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. 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 90o; 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 45o 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. 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.; 27o 44'N, 117 o 33'E, 1200���1500m; 25-V-2012, PENG & DAI leg. (2). Deposited in SNUC, FSCA. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 29-31, DOI: 10.5281/zenodo.575641, {"references":["Mader, L. (1939) Neue Coleopteren aus China. Entomologisches Nachrichtenblatt, 13 (1 - 2), 41 - 51.","Wegrzynowicz, P. (2007) Family Erotylidae Latreille, 1802. In: Lobl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera, Folume 4. Apollo Books, Stenstrup, pp. 531 - 546.","Mader, L. (1955) Neue Coleopteren aus Fukien (China). Helotidae, Languriidae, Erotylidae, Endomychidae, Coccinellidae. Koleopterologische Rundschau, 33 (1 - 6), 62 - 78.","Ye, Z. - Y. (1999) The investigation into the past and the current state of Cycas revoluta in Fujian. In: Chen, C. J. (Ed.), Biology and Conservation of Cycads. International Academic Publishers, Beijing, pp. 73 - 74.","Liu, N. & Qin, G. (2004) Notes on some species of Cycas (Cycadaceae) from China. In: Lindstrom, A. J. (Ed.), Proceedings of CYCAD 2002. Nong Nooch Tropical Garden, Bangkok, pp. 1 - 4."]}

Cycadophila (Cycadophila) torquata Skelley, Xu & Tang, new species Figs. 11 A���E 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. 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 45o 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. 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. Etymology. The species epithet, ��� torquata ���, is Latin for adorned with a necklace or collar, in reference to the prominent ridges (temples) behind the eyes. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 36-38, DOI: 10.5281/zenodo.575641

Cycadophila (Cycadophila) debaonica Xu, Tang & Skelley Figs. 2 E, 2H–I, 2L, 2P, 3C, 3D, 3I Cycadophila debaonica Xu, Tang & Skelley in Xu et al. 2015: 7 –12. 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 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 16S rRNA gene (GenBank Accession numbers KR005715 – KR00717, KY365239, KY365249, KY365251), as discussed earlier. Full descriptions and images of adults and larvae are presented in Xu et al. (2015). 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, 50–100m, 25-IV- 2015, N. S. Khang ex ♂ Cycas hoabinhensis (1; deposited at the FSCA). 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 16S 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.

Cycadophila (Cycadophila) abyssa Skelley, Xu & Tang, new species Figs. 6 A���E 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. 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 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;����� 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. Mat�� (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���1000m, 15��54'N, 109��41'E, 18-IV-2012, YIN Ziwei (1). Paratypes deposited in BMNH, FSCA, NZAC, SNUC. Etymology. The specific epithet is derived from the Greek ��� abyssos��� meaning bottomless pit, used in reference to the pit at the submental-gular juncture. 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., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 23-25, DOI: 10.5281/zenodo.575641

Key to larvae of Pharaxonotha floridana, P. kirschii and known larvae of Cycadophila Analysis of the 16S rRNA gene revealed an identical match between one specimen of a newly recognized larval morphotype collected in a male cone of Cycas petraea (D0331 in Fig. 1) and adult Cycadophila (Strobilophila) tansachai found in Cycas petraea, C. elephantipes and C. pectinata. A second sample of this larval morphotype (D0333 in Fig. 1) from a male cone of C. pectinata in Thailand, however, did not match these sampled adults. Based on this analysis this larval morphotype is placed in subgenus Strobilophila, but is not assigned to a specific species. The unmatched Strobilophila larva may correspond with Cycadophila (S.) yangi , one of two species of adults of subgenus Strobilophila found together on the cones of Cycas pectinata in Thailand [see also Remarks under description of C. (S.) yangi]. The key to larvae below includes our new morphotype of Cycadophila (Strobilophila) spp., two species of Cycadophila (Cycadophila) previously described by Xu et al. (2015) and two other larvae of Pharaxonothinae including Pharaxonotha kirschii (illustrated by B��ving & Craighead 1930) and Pharaxonotha floridana. 1. Paired tubercles t1 anterior to urogomphi on A9 without apical hook, but with a short subapical spine pointing posteriad or with apical hook and subapical spine both absent............................................................ 2 - Paired tubercles t1 anterior to urogomphi on A9 (Figs. 3 C���F, 15C���D) each with an apical hook, shaped like a bird head, pointing posteriad..................................................................... Cycadophila sp. ... 3 2. Paired tubercle t1 with a short subapical spine pointing posteriad (Figs. 3 A���B).......... Pharaxonotha floridana (Casey) - Paired tubercle t1 without a subapical spine........................................ Pharaxonotha kirschii Reitter 3. Urogomphi length measured from tubercle t3 (Fig. 3 C���D) Cycadophila (Cycadophila) debaonica Xu, Tang & Skelley - Urogomphi length measured from tubercle t3 (Fig. 3 E���F) = or>length of segment A9; tubercle t1 anterior to urogomphi with base vertical or tilting in postero-dorsal direction toward apices of urogomphi (Figs. 3 E, 15C); incisors of mandible not serrate (Figs. 3 H, 15F)...................................................................................... 4 4. Length of antennomere II ~ 2�� own width (Fig. 3 G); sclerotized dorsal plates on abdominal segments A1���A8 with 2 pairs of tubercles along posterior margin, tubercle color brown, slightly darker than dorsal plates (Fig. 3 F); turbercle t3 on A9 prominent in dorsal view, larger than t2 (Fig. 3 F)............ Cycadophila (Cycadophila) fupingensis Skelley, Tang & Xu - Length of antennomere II ~ 3�� own width (Fig. 15 E); sclerotized dorsal plates on abdominal segments A1���A8 with 5���6 pairs of tubercles along posterior margin, tubercles light-colored, contrasting with brown of dorsal plates (Fig. 15 D); turbercles t2 and t3 on A 9 in dorsal view equal in size (Figs. 15 D)...................... Cycadophila (Strobilophila) new subgenus, Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 11-12, DOI: 10.5281/zenodo.575641, {"references":["Xu, G., Tang, W., Skelley, P., Liu, N., & Rich, S. (2015) Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa, 3986 (3), 251 - 278.","Boving, A. G. & Craighead, F. C. (1930) An illustrated synopsis of the principal larval forms of the order Coleoptera. Entomologica Americana, 11 (1), 1 - 34, Pl. 28."]}

Cycadophila (Cycadophila) vittata (Arrow), new combination Figs. 12 A���F Pharaxonotha vittata Arrow 1925: 255, fig. 46; Schenkling 1928: 31; Mader 1939: 47; Ch��j�� 1964: 224; Ch��j�� 1967: 41. 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. 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. 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). 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 (Lindstr��m, unpub. data)., Published as part of Skelley, Paul, Xu, Guang, Tang, William, Lindstr��m, 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, pp. 1-63 in Zootaxa 4267 (1) on pages 38-40, DOI: 10.5281/zenodo.575641, {"references":["Arrow, G. J. (1925) Coleoptera. Clavicornia. Erotylidae, Languriidae, and Endomychidae. In: Shipley, A. E. & Scott, H. (Eds.), The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, XVI + 416 pp., 76 figs., pl., map.","Schenkling, S. (1928) Languriidae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus, Pars 100, 40 pp.","Mader, L. (1939) Neue Coleopteren aus China. Entomologisches Nachrichtenblatt, 13 (1 - 2), 41 - 51.","Chujo, M. (1964) Coleoptera from Southeast Asia (III). 20. Family Languriidae. Nature and Life in Southeast Asia, 3, 219 - 226, pl. 14.","Chujo, M. (1967) Languriid-Beetles from Thailand, Laos, Viet-Nam, S. China, Is. Hainan, Formosa and Ryukyus. Studies on the Languriid-Beetles (12). Memoirs of the Faculty of Education Kagawa University, 2 (149), 1 - 42."]}

Cycadophila nigra (Gorham), new combination (Figure 8) Thallis ? nigra Gorham, 1895: 324. Pharaxonotha nigra (Gorham); Arrow, 1925: 255; Schenkling, 1928: 31; Ch��j��, 1964: 224, pl. 14, fig. 7; Ch��j��, 1967: 41. Pharaxonotha indica Grouvelle, 1903: 127; Grouvelle, 1916: 62 (sp. bona); Schenkling, 1923: 15 (sp. bona); Arrow, 1925: 255 [syn.]; Schenkling, 1928: 31; Ch��j��, 1964: 224. Adult diagnosis. Distinguished from other Cycadophila spp. by its larger, entirely brown body, distinct supraocular stria that is half the length of the eye, weakly depressed submental-gular region, and male with a large angulate swelling near the middle of the medial mesotibial margin. Adult male description. Length 4.35���4.59 mm, width 1.65���1.76 mm. Body in dorsal view elongate, sides evenly arcuate, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color entirely dark reddish-brown; elytra lacking maculation of any kind, although lateral margins may be paler; dorsal surface distinctly punctate, surface appearing slightly shining, dorsally appearing glabrous, short procumbent hairs barely protruding out of punctures, ventrally mostly covered with short procumbent setae (Figs. 8 A���C). Head in dorsal view conical, gradually narrowed anteriorly, surface convex, distinctly punctured (Fig. 8 E); width 1.0��� 1.1 mm; dorsal interocular distance 0.63���0.68 mm, head width/dorsal interocular distance ratio 1.56��� 1.60, ventral interocular distance 0.40���0.45 mm, head width/ventral interocular distance ratio 2.33���2.50. Eye globular; bordered dorsally with strong supraocular stria at basal 1 / 2 of eye; projecting laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with a 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 about 3 / 4 pronotal width, slightly longer than head width, antennomere relative lengths approximately 13: 10: 12: 10: 10: 10: 10: 10: 13: 13: 17; 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 anteriorly rounded, 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 �� width, edge nearly straight, with fringe of hairs along dorsal base that extends to the proximal apical tooth; mola with carinulation and small spines. Labrum transverse, about 2 times 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 palp 4 -segmented, palpomere I very small; 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 palp 3 -segmented; palpomere I small, elongate; II transverse; III large, ovate. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum, where shallow circular depression is present with setose punctures. Thorax with pronotum transverse in dorsal view with marginal beads on all sides; length/width ratio 0.69���0.71, convex, nearly parallel-sided; anterior angles projecting, weakly angulate; lateral margins nearly straight, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 o; posterior margin projecting medially; with a 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���0.22. Prosternum in ventral view convex; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae; prosternal process truncate apically. Hypomeron laterally with coarse punctures, medially without longitudinal striations. Scutellar shield transverse, posterior margin clearly projected, pentagonal. Elytra in dorsal view elongate, evenly arcuate along sides, convex; length/width 3.47���3.57, greatest width near midlength; with marginal line basally, 10 complete striae of moderate puncture size, scutellary striole with 9���10 punctures; intervals of striae with distinct 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 distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora compressed laterally; tibiae shorter than femora, weakly dilated to narrow apex. Protibia not dilated apically, with fine setal fringe ventrally, lateral margin smooth, lacking stout spinules. Meso- and metatibia narrow, similar to protibia. Metatibia with angulate swelling on apical half of medial margin (Fig. 8 F). 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 adjacent to the midline (at least none were observed on available specimens); II���IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex, with row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left 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 coniform-shaped parameres. Parameres apically with long setae; length/width ratio 2.5. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of penile strut length to median lobe length 5: 1. Female similar to male except lacking swelling on inner metatibial margin; with pygidium truncate, not as well developed. Genitalia with gonocoxites triangular, gradually narrowing 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 elongate, Cshaped. Type locality. Kanara, S. India. Range. S. India, Thailand, China. Material examined. Lectotype (female) here designated of Thallis nigra Gorham with the following labels (Figs. 8 A���D): 1) [rectangular; white with yellow horizontal line; printed in black ink] Kanara, S. India; 2) [rectangular; white; printed in black ink] Andrewes Bequest B. M. 1922 ��� 221; 3) [white circle, ringed with red ink; printed in black] Type; 4) [rectangular; red; handwritten in black ink] Type Gorham; 5) [rectangular; white; hand written in black ink] nigra (Gorh.; 5) [rectangular red paper] LECTOTYPE Thallis nigra Gorham, des. P. Skelley 2014 (BMNH, examined). Paralectotype (female) here designated of Thallis nigra Gorham with the following labels: 1) [rectangular; white with yellow horizontal line; printed in black ink] Kanara, S. India; 2) [rectangular; red; handwritten in black ink] Type Gorham; 5) [rectangular; white; hand written in black ink] Thallis nigra, Gorh. 6) [rectangular yellow paper] PARALECTOTYPE Thallis nigra Gorham, des. P. Skelley 2014 (BMNH, examined). Holotype (teneral female) of Pharaxonotha indica Grouvelle with the following labels: 1) [rectangular; white with yellow horizontal line; printed in black ink] Kanara, S. India; 2) [rectangular; white; printed in black ink] Andrewes Bequest B. M. 1922 ��� 221; 3) [rectangular; white; printed in black ink] Kanara; 4) [white circle, ringed with red ink; printed in black] Type; 5) [rectangular; red; printed in black ink] Type; 6) [rectangular; white; hand written in black ink] Pharaxonota [sic] indica 5. Grouv (BMNH, examined). Other materials examined: [India:] JAMUN, 22 - 7-69, Durgapura, C.I.E. Coll. A. 3753, V 235 / 133, Pres by Com Inst Ent B M 1970 - 1, Pharaxonotha nigra (Gorh.) R. Madge det 1970 (1 M-BMNH) [Although there is a Jamun, India, we suspect Jamun is a plant on which it was collected, and Durgapura is a locality name of multiple localities in India]. Central India, Pachmarhi, Sept. 1970, T. R. Nathan leg. (1 F-FSCA). CHINA: Guangxi: [Napo Co.]: Dingye, ex ♂ cone Cycas debaoensis, N 23 �� 24 ' 26 ���, E 106 ��02��� 27 ���, 22 -V- 2006, W. Tang, # 1 (21); # 2 (1). These Chinese specimens are deposited at FSCA, IZCAS, and NZAC. Remarks. Smaller females of C. nigra are difficult to distinguish from unicolorous C. yunnanensis. Gorham did not designate a holotype from the specimens he studied. The specimen here designated as the lectotype is Gorham���s darker and more intact specimen. It is unfortunate he only had females to study because sexual dimorphisms are important for species delineation. With the types of C. nigra being from southern India, there is reason to question whether the Chinese specimens here considered to be C. nigra are truly conspecific. More material is needed from southern India to better establish character states (both morphological and molecular) of those populations in comparison with those from China. For now, they are considered the same species. In Cycas debaoensis adults of this species have only been detected in small numbers on male cones of the Dingye population. The relative abundance of this species among Cycadophila -inhabiting cones at Dingye ranges 1.5��� 24.1 % (mean = 12.8 %, n = 2 cones). No larval morphotypes have been observed in cones that may correspond to this species. Their infrequent occurrence on cones and 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., Published as part of Xu, Guang, Tang, William, Skelley, Paul, Liu, Nian & Rich, Stephen, 2015, Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia, pp. 251-278 in Zootaxa 3986 (3) on pages 267-270, DOI: 10.11646/zootaxa.3986.3.1, http://zenodo.org/record/240311, {"references":["Gorham, H. S. (1895) List of the Coleoptera in the collection of H. E. Andrewes Esq. from India and Burma, with descriptions of new species and notes. Families: Malacodermata- Erotylidae- Endomychidae. Annales de la Societe Entomologique de Belgique, 39, 293 - 330.","Arrow, G. J. (1925) Coleoptera. Clavicornia. Erotylidae, Languriidae, and Endomychidae. In: Shipley, A. E. & Scott, H. (Eds.), The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, pp. I - XVI + 1 - 416.","Schenkling, S. (1928) Languriidae. Coleopterorum Catalogus, 100, 1 - 40.","Chujo, M. (1964). Coleoptera from Southeast Asia (III). 20. Family Languriidae. Nature and Life in Southeast Asia, 3, 219 - 226.","Chujo, M. (1967) Languriid-Beetles from Thailand, Laos, Viet-Nam, S. China, Is. Hainan, Formosa and Ryukyus. Studies on the Languriid-Beetles (12). Memoirs of the Faculty of Education Kagawa University, 2, 1 - 42.","Grouvelle, A. (1903) Coleopteres de l'Inde recoltes par MM. H. - L. Andrewes et T. - R. - D. Bell. Annales de la Societe Entomologique de France, 72, 125 - 127.","Grouvelle, A. (1916) Descriptions d'especes nouvelles de Cryptophagidae. Memoires Entomologiques. Etudes sur les Coleopteres, 1, 30 - 79.","Schenkling, S. (1923) Cryptophagidae. Coleopterorum Catalogus, 76, 1 - 92."]}

Cycadophila yunnanensis (Grouvelle), new combination (Figures 9���10) Pharaxonotha yunnanensis Grouvelle, 1916: 61; Schenkling, 1923: 15; Mader, 1926: 721; Schenkling, 1928: 31; Wegrzynowicz, 2007: 536. Adult diagnosis. Distinguished from other Cycadophila spp. by its smaller body with or without color pattern, distinct supraocular stria that is less than 1 / 3 length of the eye, weakly depressed submental-gular region, and male with a small rounded swelling near the middle of the medial metatibial margin. Adult male description. Length 3.12���3.88 mm, width 1.29���1.65 mm. Body in dorsal view elongate, sides nearly straight and parallel sided, greatest width at middle of elytra; in lateral view convex dorsally. General body and appendage color dark reddish-brown, pronotum occasionally with orange-brown sides; elytra of type specimens entirely dark brown (Fig. 9) or with light central stripe (see variation in Figs. 10 A���C); dorsal surface distinctly punctate, surface appearing shiny, glabrous, short procumbent hairs protruding out of punctures, ventrally mostly covered with short procumbent setae (Figs. 9 C���F). Head in dorsal view conical, gradually narrowed anteriorly, surface convex, distinctly punctured (Figs. 9 C, E); width 0.8���0.9 mm; dorsal interocular distance 0.48���0.55 mm, head width/dorsal interocular distance ratio 1.68, ventral interocular distance 0.40���0.45 mm, head width/ventral interocular distance ratio 2.00��� 2.06. Eye globular; bordered dorsally with short supraocular stria at basal 1 / 3 of eye; projecting laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with a 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 about 3 / 4 pronotal width, slightly more than head width, antennomere relative lengths approximately 16: 13: 14: 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. Mandible with 3 apical teeth; prostheca extending from mola halfway to apical teeth, length,about �� width, edge nearly straight, with fringe of hairs along dorsal base that extends to the proximal apical tooth; mola with carinulation and small spines. Labrum transverse, about 2 times 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 palp 4 -segmented, palpomere I very small; 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 palp 3 -segmented; palpomere I small, elongate; II transverse; III large, ovate. Mentum and submentum with moderate punctation and setation, setae short. Gular area smooth, without setae or punctation, except near suture with submentum, where shallow depression present with setose punctures. Thorax with pronotum transverse in dorsal view; with marginal beads on all sides; length/width ratio 0.64��� 0.70, convex, nearly parallel-sided; anterior angles projecting, weakly angulate; lateral margins nearly straight, with thin bead, bead bearing row of setose punctures; posterior angles sharp, nearly 90 o; posterior margin projecting medially; with a narrow, short longitudinal furrow posterolaterally, beginning at pore on posterior margin, extending forward and ending abruptly, length of furrows relative to pronotal length 0.14���0.18. Prosternum in ventral view convex; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae; prosternal process truncate apically. Hypomeron laterally with coarse punctures, medially without longitudinal striations. Scutellar shield transverse, posterior margin clearly projected, pentagonal. Elytra in dorsal view elongate, nearly parallel sided, length/width 3.29���3.45, greatest width near midlength; with marginal line basally, 10 complete striae of moderate puncture size, scutellary striole with 8���10 punctures; intervals of striae with distinct 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 distinct. Metaventrite long, convex laterally, slightly impressed medially, metathoracic discrimen extending slightly over half metaventrite length. Legs narrow, similar in length; procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora compressed laterally; tibiae shorter than femora, weakly dilated to narrow apex. Protibia not dilated apically, with fine setal fringe ventrally, lateral margin smooth, lacking stout spinules. Meso- and metatibiae narrow, similar to protibia. Metatibia with slight rounded swelling on apical half of medial margin (Figs. 10 B���C). 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 adjacent to the midline (at least none were observed on available specimens); II���IV subequal in length; V slightly longer than IV with lateral margins converging posteriorly to a rounded apex, with row of short appressed, densely arranged setae. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left and resting on side (Fig. 9 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 coniform-shaped parameres. Parameres apically with long setae; length/width ratio 2.0. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of penile strut length to median lobe length 5: 1. Female similar to male except with metatibia without slight rounded swelling on apical half of medial margin (Fig. 10 A), pygidium truncate, not as well developed. Genitalia with gonocoxites triangular, gradually narrowing 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 posteriorly expanded. Spermatheca elongated, C-shaped. Variation. The original syntypes of C. yunnanensis (Fig. 9 A) are mostly teneral, with two dark adults. The teneral specimens exhibit a vague central stripe on each elytra (Fig. 10 B). Series of specimens from independent collecting events show greater variation in the distinctness of this central elytral stripe as well as the presence of lateral stripes (Fig. 10 C). Elytra of these series vary from entirely dark brown to sharply striped with intermediate color morphs displaying varying degrees in size and distinctness of these stripes, as well as some in which the stripes are reduced to a single humeral and apical spot. Type locality. Yunnan [Coll. Grouvelle-Paris]. Range. Known from Guangxi and Yunnan provinces, China and Manipur state, India. Material examined. Six syntypes with the following labels (Fig. 9 B): 1) [rectangular, red, hand written in black ink] Yunan Mission; 2) [rectangular, pale red, printed in black ink] Type; 3) [rectangular, white, hand written in black ink] Pharaxonota [Pharaxonotha] yunnanensis Grouv. (Grouvelle collection - BMNH, examined). The sixth specimen, a male (Figs. 9 A, C���G) is here designated as the lectotype and placed on a separate card labeled as lectotype. The remaining specimens are here designated as paralectotypes. A red lectotype and yellow paralectotype labels stating ��� Pharaxonotha yunnanensis Grouvelle, des. P. Skelley 2014 ��� have also been placed on the pin with these specimens. Other materials considered C. yunnanensis, that show much color variation as discussed above, include: CHINA: Guangxi: [Debao Co. ]: Fuping, ex ♂ cone Cycas debaoensis, N 23 �� 29 ��� 50 ��� E 106 �� 12 ��� 87 ���, V- 2001, W. Lu (9); N 23 �� 29.643 ', E 106 �� 12.915 ', 21 -V- 2004, W. Tang, # 1 (542); N 23 �� 29.663 ', E 106 �� 12.903 ', 21 -V- 2004, W. Tang, # 7 (13); 21 -V- 2004, W. Tang, # 31 (31); N 23 �� 29.595 ', E 106 �� 12.944 ', 21 -V- 2004, W. Tang, # 210 (78); N 23 �� 29.624 ', E 106 �� 12.980 ', 21 -V- 2004, W. Tang, # 331 (30); N 23 �� 29.643 ', E 106 �� 12.914 ', 26 -V- 2006, W. Tang, # 1 (31); N 23 �� 29.601 ', E 106 �� 12.862 ', 26 -V- 2006, W. Tang, # 2 (37); N 23 �� 29.597 ', E 106 �� 12.863 ', 26 -V- 2006, W. Tang, # 4 (11); 24 -V- 2008, W. Tang (75); [Napo Co.]: Dingye, N 23 �� 24 ��� 26 ��� E 106 ��01��� 27 ���, 22 -V- 2004, W. Tang, # 1 (56); # 2 (20); India: Manipur: Nongmaiching, ex Cycas pectinata ♀ cone, 2007, J. Singh & R. Singh (2). These are deposited at ANIC, BMNH, FSCA, IZCAS, MNHN, NZAC, USNM. Remarks. Specimens placed here show color pattern variation from solid color to striped elytra (Fig. 10), with intermediate color morphs. Molecular analysis of the 16 S rRNA gene for 25 individuals of the C. yunnanensis morphotype from the Fuping population reveals three variants of this gene (Fig. 11). At least two of these genotypes are found in the Napo population and one as far away as Manipur, India on another species of Cycas (data not shown). This color variation is found within at least two of these genotypes. Although morphological analysis has not yielded any data to support more than one species being present, the genetic variation is relatively high for this species and there may be cryptic species present. Dissection of adults confirms cycad pollen in their guts. Adults are found in small numbers in male cones of Cycas debaoensis at the early stage of elongation and pollen shed and they account for a small percentage of the adult Cycadophila present at this stage (range 3.3��� 30.3 %, mean = 10.9 %, n = 5 cones), but numbers increase up to hundreds in cones that have completed pollen shedding and C. yunnanensis usually makes up the highest proportion of adult Cycadophila at this stage (range 41.1 ���100.0%, mean = 75.3 %, n = 7 cones). No larval morphotype has been found in Cycas debaoensis cones which may correspond with C. yunnanensis, and this species may be also be an opportunistic visitor that reproduces and feeds on alternative hosts., Published as part of Xu, Guang, Tang, William, Skelley, Paul, Liu, Nian & Rich, Stephen, 2015, Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia, pp. 251-278 in Zootaxa 3986 (3) on pages 270-275, DOI: 10.11646/zootaxa.3986.3.1, http://zenodo.org/record/240311, {"references":["Grouvelle, A. (1916) Descriptions d'especes nouvelles de Cryptophagidae. Memoires Entomologiques. Etudes sur les Coleopteres, 1, 30 - 79.","Schenkling, S. (1923) Cryptophagidae. Coleopterorum Catalogus, 76, 1 - 92.","Mader, L. (1926) Clavicornia [part]. In: Winkler, A. (Ed.), Catalogus Coleopterorum regionis palearcticae. Pars 6. A. Winkler, Wien, pp. 625 - 752.","Schenkling, S. (1928) Languriidae. Coleopterorum Catalogus, 100, 1 - 40.","Wegrzynowicz, P. (2007) Family Erotylidae Latreille, 1802. In: Lobl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 4. Apollo Books, Stenstrup, pp. 531 - 546."]}

Key to species of larval Cycadophila occurring with Cycas debaoensis 1 Urogomphi shorter than segment A 9; hooks anterior to urogomphi (Figs. 4 C–D) with base tilting in antero-dorsal direction away from apices of urogomphi, distal portion of hooks shaped like a bird’s bill, with tip of bill not sharply curved..................................................................................... Cycadophila debaonica, n. sp. - Urogomphi longer than segment A 9; hooks anterior to urogomphi (Figs. 7 C–D) with base tilting in dorsal to postero-dorsal direction towards apices of urogomphi, distal portion of hooks shaped like a bird’s bill, with tip of bill sharply curved................................................................................. Cycadophila fupingensis, n. sp.

Xu, Guang, Tang, William, Skelley, Paul, Liu, Nian, Rich, Stephen (2015): Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia. Zootaxa 3986 (3): 251-278, DOI: http://dx.doi.org/10.11646/zootaxa.3986.3.1

Genus Cycadophila Xu, Tang & Skelley, new genus Type species. Cycadophila debaonica n. sp., here designated. Diagnosis. Cycadophila is distinguished from other currently known cycad-associated genera of Erotylidae by the presence of a transverse occipital ridge; supraocular striae present; having a remnant of the submental-gular suture visible to variously depressed; lack of abdominal calli; male genitalia with median lobe and tegmen twisted like a corkscrew, and spiculum gastrale asymmetrical; wings with anal cell present. Description. Length 3.12���5.27 mm, width 1.29���2.31 mm. Body form elongate, not strongly parallel sided; dorsoventrally flattened, weakly convex; widest anterior to or at middle of elytra; color pale to dark reddish-brown with or without variously sized dark macula on elytra; dorsal punctation fine, evenly distributed. 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, about 1 / 3 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 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. Scutellum broadly pentagonal. Elytra finely setose, setae indistinct; base of elytra with marginal bead. Prosternum mostly flat, depressed posterior of coxa; procoxal cavities narrowly open; 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. Etymology. The name Cycadophila is formed from the words ���cycad��� and ���philia���, ancient Greek for affection. Gender feminine. Remarks. With the addition of Cycadophila there are now 6 genera in Pharaxonothinae. However, an understanding of the relationship of Cycadophila within Pharaxonothinae, particularly with respect to Pharaxonotha, requires further studies using morphological characters and multilocus sequence analysis. Furthermore, a phylogenetic comparison of pharaxonothines and their food plants may sharpen our understanding of their co-evolutionary relationships. All specimens of Cycadophila with biological data were collected from male cones of Asian cycads, Cycas spp. Tang et al. (1999) divided ��� Xenocryptus ��� (now Cycadophila) into different groups (XA and XB, for ��� Xenocryptus A��� and ��� Xenocryptus B���) based on an initial assessment of morphology. In field observations, Tang (see below) noted members of the XA type (C. debaonica and C. fupingensis) appear to reproduce in the male cones with fresh pollen. Specimens of the XB type (C. nigra and C. yunnanensis) are abundant on cones that have finished shedding their pollen and may be saprophytes feeding on old rotting cones, pollen and fungi. There are presently four species herein placed in the genus Cycadophila that occur on cones of C. debaoensis. Other Asian species previously placed in ��� Pharaxonotha ��� and additional new species of Cycadophila that occur on other species of cycads will be treated in a separate future publication (Xu et al. in prep) reviewing all known members of the genus. We here focus only on the species of Cycadophila known to occur on C. debaoensis., Published as part of Xu, Guang, Tang, William, Skelley, Paul, Liu, Nian & Rich, Stephen, 2015, Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia, pp. 251-278 in Zootaxa 3986 (3) on pages 255-256, DOI: 10.11646/zootaxa.3986.3.1, http://zenodo.org/record/240311, {"references":["Tang, W., Oberprieler, R. & Yang, S. L. (1999) Beetles (Coleoptera) in Asian Cycas: diversity, evolutionary patterns, and implications for taxonomy. In: Chen, C. J. (Ed.), Biology and Conservation of Cycads. International Academic Publishers, Beijing, pp. 280 - 297."]}

Cycadophila debaonica Xu, Tang & Skelley, new species (Figures 2–4) Adult diagnosis. Distinguished from other Cycadophila spp. by the long supraocular striae, head width/ventral interocular distance ratio Adult male description. Length 3.86–5.27 mm, width 1.70–2.31 mm. Body in dorsal view elongate-oval, greatest width at middle of elytra; in lateral view convex dorsally. General body color reddish-brown (Figs. 2 A–D), head darker than pronotum and lateral halves of elytra; elytra with large central black maculation near medial margin 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. Head in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, finely, sparsely punctured (Figs. 3 A, C); width 0.89–1.16 mm; dorsal interocular distance 0.48–0.70 mm, head width/dorsal interocular distance ratio 1.42–1.91, ventral interocular distance 0.30–0.41 mm, head width/ventral interocular distance ratio 2.92–3.13. 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 approximately 18: 12: 13: 10: 10: 10: 10: 10: 14: 14: 21; 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. 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 times 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 palp 4 - segmented, palpomere I very small; II, III and IV longer than width; 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 3 -segmented; palpomere I small, elongate; II transverse; III large, 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 fovea are present bearing dense setose punctures (Fig. 3 D), fovea dark brown. Thorax with pronotum transverse in dorsal view with marginal beads on all sides; length/width ratio 0.60–0.69, 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 0.16–0.19. Prosternum in ventral view convex; 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 2.7–3.3, greatest width near midlength; with marginal line basally, 10 complete striae of moderate puncture size, 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. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; 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 (Fig. 3 E). 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 (Fig. 2 E). 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-shaped parameres. Parameres apically with long setae; length/width ratio 2.6. 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 protibia less dilated apically, mesotibia slightly less robust than in male, 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 tibia 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. Larval description. Mature individuals 6.5 mm; body elongate, slightly curved ventrally, in dorsal view more or less parallel sided and narrowing slightly anteriorly and posteriorly, cylindrical, lightly pigmented except for head, legs and sections of tergum (Figs. 4 A–E). Head sclerotized, light brown with faint pattern of longitudinally elongate reticulation, smooth without ornamentation, protracted, prognathous and slightly flattened; epicranial stem absent, frontal arms lyriform and contiguous basally; median epicranial suture distinct and dark, extending nearly the length of frontal arms; 5 stemmata on each side; antenna length just less than half of dorsal length of head at midline, 3 -segmented, antennomere 1 slightly shorter than wide, 2 almost twice as long as 1, 3 slightly shorter than 1, sensorium located on ventral apex of antennomere 2, conical, with convex sides, length about 1 / 2 length of antennomere 3, width slightly more than 1 / 2 width of antennomere 3; frontoclypeal suture absent; labrum free. Mandibles symmetrical, incisor cleft with dorsal and ventral prongs of nearly equal length, dorsal prong serrate and ventral prong smooth; mola well-developed, asperate with asperites forming transverse rows; accessory ventral process present; prostheca hyaline, with two edges, dorsal edge broadly rounded, ventral edge broad basally and obtusely angulate (Fig. 4 F). Ventral mouthparts retracted, with well-developed maxillary articulating area; maxilla with transverse cardo, elongate stipes, 3 -segmented palp, terminal palpomere 2 times 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 1.5 times width of basal palpomere (Fig. 4 G), apical palpomere length 1.5 times width. Hypostomal rods diverging; ventral epicranial ridges weakly developed. Thorax and abdomen with central ½ of T 1, anterior ½ of T 2–3 and A 1 –A 7, anterior ¾ of A 8, and all of A 9 pigmented light brown dorsally, sclerotized; sclerotized regions of T 1–3 bisected by narrow light-colored midsagittal line; brown areas of all thoracic and abdominal segments, except for their margins, more or less randomly covered with granules; T 2–3, A 1–8 with straight row of 3 or more pairs of setiferous tubercules along posterior edge of granule field; A 9 tergum with pair of urogomphi; base of urogomphi covered with granules, dorsal and lateral margins 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. 4 C–D) with hook at apex, bases of this pair angled in antero-dorsal direction, apical hook shaped like a bird head and neck with bill pointing posteriad (Figs. 4 C–D), top of “bird head” with rounded prominence and single seta, setal length slightly less than length of tubercle; “bird beak” length about 1 times its vertical width, without sharply curved tip; urogomphi length from basal tubercle t 3 to apex 1 times coxal width; meso- and metacoxae separated by> 2 times coxal width (Fig. 4 E). Spiracles raised, annular-biforous. Type locality. China, Guangxi Province, Debao County, Fuping village, N 23 ° 29.624 ', E 106 ° 12.980 '. 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 (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] CHINA, Guangxi, [Debao,] Fuping, ex ♂ cone Cycas debaoensis, N 23 ° 29.624 ', E 106 ° 12.980 ', 21 -V- 2004, W. Tang, # 331; 2) [rectangular; red; printed in black ink] HOLOTYPE ♂ Cycadophila debaonica G. Xu, W. Tang & P. Skelley 2015. Deposited in the FSCA. Allotype (FSCA) and 1683 adult + 329 larval paratypes: CHINA: Guangxi: [Debao Co. ]: Fuping, ex ♂ cone Cycas debaoensis, N 23 ° 29 ’ 50 ” E 106 ° 12 ’ 87 ”, V- 2001, W. Lu (241); N 23 ° 29.643 ', E 106 ° 12.915 ', 21 -V- 2004, W. Tang, # 1 (146); N 23 ° 29.663 ', E 106 ° 12.903 ', 21 -V- 2004, W. Tang, # 7 (92); N 23 ° 29.655 ', E 106 ° 12.867 ', 21 -V- 2004, W. Tang, # 31 (97 larvae); N 23 ° 29.595 ', E 106 ° 12.944 ', 21 - V- 2004, W. Tang, # 210 (31 larvae); N 23 ° 29.624 ', E 106 ° 12.980 ', 21 -V- 2004, W. Tang, # 331 (holotype & allotype, FSCA, 535); N 23 ° 29.643 ', E 106 ° 12.914 ', 26 -V- 2006, W. Tang, # 1 (571); N 23 ° 29.601 ', E 106 ° 12.862 ', 26 -V- 2006, W. Tang, # 2 (50 + 85 larvae); N 23 ° 29.669 ', E 106 ° 12.909 ', 26 -V- 2006, W. Tang, # 4 (1 + 8 larvae); 24 -V- 2008, W. Tang (2 + 108 larvae); [Napo Co.]: Dingye, N 23 ° 24 ’ 26 ” E 106 °01’ 27 ”, 22 -V- 2004, W. Tang, # 1 (5); # 2 (38). Paratypes deposited at ANIC, BMNH, FSCA, IZCAS, MNHN, NZAC, USNM. Etymology. Named for the county of collection. Remarks. The only known host of this beetle is Cycas debaoensis. This cycad occurs in small, relict populations (Tang et al. 2004; Xie et al. 2005) and this beetle species has only been collected from the male cones of this host at two localities. Determination and description of larvae were based on large numbers found associated with adults in C. debaoensis male cones and partial 16 S rRNA gene sequences which were identical to the adults. Field observations of the larvae indicate they feed and develop on the male cones of this species in large numbers, feeding on sporophyll tissue. The adults remain on the male cones by the hundreds and dissection of adults confirm cycad pollen in their guts. They typically account for the highest percentage of adult Cycadophila beetles on male cones sampled in the early stage of cone elongation and pollen shed (range 57.6–89.3 %, mean = 79.8 %, n = 5 cones), but their abundance drops in cones that have nearly completed pollen shedding (range 0–55.6 %, mean = 13.6 %, n = 7 cones). Adults of other species of Cycadophila of the “XB” type (Tang et al. 1999) and identified as C. nigra and C. yunnanensis (see below) occur sympatrically within the same cones of C. debaoensis, these being more abundant on male cones that have nearly completed pollen shedding (range = 41.1–100 %, mean = 80.7 %, n = 7 cones). Larvae of the XB type have not been detected in these cones.

Cycadophila fupingensis Skelley, Tang & Xu, new species (Figures 5���7) Adult diagnosis. Distinguished from other Cycadophila spp. by the short supraocular striae, narrow protibia lacking stout apical setae, fine pronotal punctation, and having a small elytral maculation. Adult male description. Length 3.53���3.76 mm, width 1.47���1.59 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 reddish-brown, elytra with central maculation along suture occupying medial half, not reaching base or apex; dorsal surface distinctly punctate, shining, 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 (Figs. 5 A���D). Head in dorsal view conical, gradually narrowed anteriorly, surface flat to slightly convex, distinctly punctured (Fig. 6 A); width 0.9 ���1.0 mm; dorsal interocular distance 0.50���0.53 mm, head width/dorsal interocular distance ratio 1.75���1.81, ventral interocular distance 0.33���0.35 mm, head width/ventral interocular distance ratio 2.69���2.71. Eyes globular; bordered dorsally with short supraocular striae at basal 1 / 3 of eye; projecting laterally, with large black facets and short, inconspicuous interfacetal setae. Frons with a 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 about 3 / 4 pronotal width, slightly longer than head width, antennomere relative lengths approximately 15: 11: 13: 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. Mandible with 3 apical teeth; prostheca extending from mola halfway to apical teeth, length,about �� width, edge nearly straight, with fringe of hairs along edge that extends to the proximal apical tooth; mola with carinulation and small spines. Labrum transverse, about 2 times broader than long; anterior margin fringed with long bristles. 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 palp 4 -segmented, palpomere I very small; 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 shallow lateral pockets; labial palp 3 -segmented; palpomere I small, elongate; II transverse; III large, 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 vague discolored depression present, marked by last set of submental punctures (Fig. 6 C). Thorax with pronotum transverse in dorsal view with marginal bead on all sides; length/width ratio 0.65���0.68, convex, sides arcuate; anterior angles rounded; lateral margin with thin marginal bead, bead bearing row of minutely setose punctures; posterior angles exteriorly angulate; posterior margin projecting medially; with a 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. Prosternum in ventral view convex; anterior margin slightly emarginate, finely denticulate with row of long, anteriorly directed setae; posterior prosternal process truncate and convex (Fig. 6 B). Hypomeron laterally with few punctures, medially without longitudinal striations. Scutellar shield transverse, posterior margin rounded, somewhat pentagonal. Elytra in dorsal view elongate, evenly arcuate along sides, convex; length/width 3.33���3.36, greatest width near midlength; with marginal line basally, 10 complete striae of moderate puncture size, scutellary striole with 10���12 punctures along anterior 1 / 4; intervals of striae with fine, distinct 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 distinct. Metaventrite long, convex laterally, metathoracic discrimen extending slightly over half metaventrite length, slightly impressed medially. Legs narrow, similar in length; procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora compressed laterally; tibiae shorter than femora, gradually dilated to narrow apex. Protibia narrow, weakly dilated toward apex, with apical fringe of small, hair-like spinules ventrally, lateral margin lacking spinules (Fig. 6 D); width at apex almost equal to length of tarsomere V. Meso- and metatibiae narrowed, obliquely angled at lateral 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 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. Male genitalia (tegmen and median lobe of aedeagus) twisted approximately 45 o toward the left and resting on side (Fig. 5 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 coniform-shaped parameres. Parameres apically with long setae; length/width ratio 2.6. Aedeagus with median lobe short, slender, highly sclerotized, apex long and acuminate. Ratio of penile strut length to median lobe length 5: 1. Female similar to male; except with protibia width at apex distinctly less than length of tarsomere V, pygidium truncate, not as well developed. Genitalia with gonocoxites triangular, gradually narrowing 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 elongate, Cshaped. Variation. There is little notable variation in the materials studied. The elytral maculation in some is notably smaller than others, but these may be teneral specimens where the dark mark is not fully developed. Larval description. Mature individuals up to 5.5 mm; body elongate, in dorsal view more or less parallelsided and narrowing slightly anteriorly and posteriorly, cylindrical, lightly pigmented except for head, legs and sections of tergum (Figs. 7 A���E). Head sclerotized, light brown with faint pattern of darker brown longitudinal streaks, smooth without ornamentation, protracted, prognathous and slightly flattened; epicranial stem absent, frontal arms lyriform and contiguous basally; median epicranial suture distinct and dark, extending over half the length of the frontal arms; 5 stemmata on each side; antenna length less than half of dorsal length of head at midline, 3 -segmented, antennomere 1 slightly shorter than wide, 2 almost twice as long as 1, 3 slightly longer than 1, sensorium located on ventral apex of antennomere 2, conical, with convex sides, length about 1 / 4 length of antennomere 3, width slightly more than 1 / 2 width of antennomere 3; frontoclypeal suture absent; labrum free. Mandibles symmetrical, incisor cleft with dorsal and ventral prongs of nearly equal length, both prongs smooth; mola well-developed, asperate with asperites forming transverse rows; accessory ventral process present; prostheca hyaline, with two edges, dorsal edge broadly rounded, ventral edge broad basally, triangular with angulate apex (Fig. 7 F). Ventral mouthparts retracted, with well-developed maxillary articulating area; maxilla with transverse cardo, elongate stipes, 3 -segmented palp, terminal palpomere slightly less than 3 times 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 1.5 times width of basal palpomere (Fig. 7 G), apical palpomere length 1.5 times its own width. Hypostomal rods diverging; ventral epicranial ridges weakly developed. Thorax and abdomen with central �� of T 1, anterior �� of T 2���3 and A 1 ���A 7, anterior �� of A 8, and all of A 9 pigmented brown dorsally, sclerotized; sclerotized regions of T 1���3 and A 1���8 distinctly bisected by light-colored midsagittal line; brown areas, except for their margins, more or less randomly covered with granules; T 2���3 with 3 large tubercles along each lateral margin of sclerotized area; A 1���8 with 2 pairs of large tubercules along posterior posterior margin; tubercles setiferous, but setae often broken off; A 9 tergum with pair of urogomphi; base of urogomphi may or may not be covered with granules, dorsal and lateral sides of base with 3 pairs of prominent setiferous tubercles, but setae may be broken off; in dorsal view tubercles form approximately semicircular pattern, pair of tubercles closest to midline just anterior to urogomphi (labeled t 1 in Figs. 7 C���D) with hook at apex, bases of this pair angled vertically to postero-dorsally in direction, apical hook shaped like a bird head and neck with bill pointing posteriad (Figs. 7 C), top of ���bird head��� with a prominence and a single setae, but setae often broken off; ���bird beak��� length about 2 times vertical width, with sharply curved tip; urogomphi length from basal tubercle t 3 to apex> 1 times length of remainder of segment; apical 1 / 3 of length smooth without tubercles. Legs 5 -segmented; coxa moderately widely separated, procoxae separated by> 1 times coxal width; meso- and metacoxa separated by> 2 times coxal width (Fig. 7 E). Spiracles raised, annular-biforous. Type locality. China, Guangxi Province, Debao County, Fuping village, N 23 �� 29.624 ', E 106 �� 12.980 '. 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 (by designation) male with the following labels: 1) [rectangular; white; printed in black ink] CHINA, Guangxi, [Debao,] Fuping, ex ♂ cone Cycas debaoensis, N 23 �� 29.624 ', E 106 �� 12.980 ', 21 -V- 2004, W. Tang, # 331; 2) [rectangular; red; printed in black ink] HOLOTYPE ♂ Cycadophila fupingensis G. Xu, W. Tang & P. Skelley 2015. Deposited in the FSCA. Allotype and 423 adult + 79 larval paratypes: CHINA: Guangxi: [Debao Co. ]: Fuping, ex ♂ cone Cycas debaoensis, N 23 �� 29 ��� 50 ��� E 106 �� 12 ��� 87 ���, V- 2001, W. Lu (20); N 23 �� 29.643 ', E 106 �� 12.915 ', 21 -V- 2004, W. Tang, # 1 (246); N 23 �� 29.663 ', E 106 �� 12.903 ', 21 -V- 2004, W. Tang, # 7 (8); N 23 �� 29.655 ', E 106 �� 12.867 ', 21 -V- 2004, W. Tang, # 31 (20 larvae); N 23 �� 29.655 ', E 106 �� 12.867 ', 21 -V- 2004, W. Tang, # 210 (2); N 23 �� 29.624 ', E 106 �� 12.980 ', 21 -V- 2004, W. Tang, # 331 (holotype & allotype, FSCA, 76); N 23 �� 29.643 ', E 106 �� 12.914 ', 26 -V- 2006, W. Tang, # 1 (53); N 23 �� 29.601 ', E 106 �� 12.862 ', 26 -V- 2006, W. Tang, # 2 (3 + 46 larvae); N 23 �� 29.669 ', E 106 �� 12.909 ', 26 -V- 2006, W. Tang, # 4 (10 larvae); 24 -V- 2008, W. Tang (3 + 3 larvae); [Napo Co.]: Dingye, N 23 �� 24 ��� 26 ��� E 106 ��01��� 27 ���, 22 -V- 2004, W. Tang, # 1 (5); # 2 (7). Paratypes deposited at ANIC, BMNH, FSCA, IZCAS, MNHN, NZAC, USNM. Etymology. Named for the type locality near the village of Fuping. Remarks. The only known host of this beetle is Cycas debaoensis and the species has only been collected from the male cones of this host at two localities. Determination and description of larvae were based on large numbers found associated with adults in C. debaoensis male cones and partial 16 S rRNA gene sequences which were identical to the adults. Field observations of the larvae indicate they feed and develop on the male cones of this species in moderate numbers, feeding on sporophyll tissue. The adults remain on the male cones in small numbers or up to hundreds and dissection of adults confirm cycad pollen in their guts. Adults and larvae occur sympatrically in the same cones with Cycadophila debaonica. The adults and larvae of C. debaonica usually outnumber those of C. fupingensis, and C. fupingensis typically accounts for a small percentage of adult Cycadophila on male cones sampled in the early stage of cone elongation and pollen shed (range 7.1���12.1 %, mean = 9.1 %, n = 5 cones). The relative abundance of adults remains about the same in cones that have nearly completed pollen shedding (range 0���26.1 %, mean = 7.5 %, n = 7 cones)., Published as part of Xu, Guang, Tang, William, Skelley, Paul, Liu, Nian & Rich, Stephen, 2015, Cycadophila, a new genus (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting Cycas debaoensis (Cycadaceae) in Asia, pp. 251-278 in Zootaxa 3986 (3) on pages 262-267, DOI: 10.11646/zootaxa.3986.3.1, http://zenodo.org/record/240311, {"references":["Xie, J., Jian, S. & Liu, N. (2005) Genetic variation in the endemic plant Cycas debaoensis on the basis of the ISSR analysis. Australian Journal of Botany, 53, 141 - 145.","Fang, S. (2009) Study on Conservated Biology of Cycas debaoensis. Sun Yat-Sen University, Guangzhou, 113 pp. [Ph. D. Dissertation]"]}