Pharaxonotha kirschii Reitter

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, Insecta Mundi 2022 (917), pp. 1-41 : 33-38

Pharaxonotha kirschii Reitter

Figures 1A , 2A, 2D–E , 12A–K

Pharaxonotha kirschii Reitter 1875: 44 ~ Sharp 1900: 598; Chittenden 1911: 8–13; Hinton 1945: 189–192; Pakaluk 1988: 450. Pharaxonotha kirschi ~ emendation of various authors.

Thallisella condradti Gorham 1898: 249 ~ Champion 1904: 36 [synonymy].

Diagnosis. A member of the kirschii group, this species is diagnosed by the uniformly dark coloration of both sexes, an enlarged antennomere XI compared to IX, the lateral pronotal marginal bead of many is weakly angulate near the middle, and the male abdominal ventrite apical tooth is more pronounced. Pharaxonotha kirschii is widespread from the southern US to Panama, is not associated with cycads. It is known as the “Mexican grain beetle”, a pest in stored plant products which has been intercepted in many countries.

Description. Length 3.37–4.11 mm, width 1.22–1.56 mm. General body color ( Fig. 12A–C ) dark brown to black. Dorsal surface glossy, with very short setae in puncture.

Head. Not broad, width = 0.62–0.67× pronotal width ( Fig. 12E–F ); in lateral view robust, clypeus to base of head dorsally convex; in dorsal view conical, gradually narrowed anteriorly, surface convex, coarsely punctured, average distance between closest punctures 2–3× width of puncture; head width 0.72–0.86 mm; dorsal interocular distance 0.51–0.60 mm, head width/dorsal interocular distance ratio 1.36–1.42, ventral interocular distance 0.41–0.51 mm, head width/ventral interocular distance ratio 1.68–1.79. Eye with large black facets, similar diameter of head punctures; head posterior of eye with a small tooth (temple) in dorsal profile. Antennal length slightly shorter than pronotal width, 1.4× head width; antennomere I (scape) fairly large, slightly elongate; antennomere II same length as III; IV circular; V–VII same length as IV, gradually becoming wider with VIII weakly transverse and flattened apically; club fairly large, IX and X similar in length; XI enlarged, 1.5× longer than X, globular with rounded apex. Clypeus weakly concave anteriorly, moderately punctate. Transverse occipital line [vertexal line] distinct nearly from eye to eye. Mentum and submentum coarsely punctured, ½–¾× 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 0.69–0.80; with distinct marginal carina laterally and basally, anteriorly with fine marginal carina medially; dorsally flattened; anterior angles sharply rounded, projecting forward; posterior angles developed, with small denticle at angle; lateral margin weakly angulate 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 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 smooth, with few punctures. Scutellar shield distinctly transverse, posterior margin weakly roundly pentagonal. Elytra in dorsal view elongate, flattened dorsally; length/width 1.72–1.90, greatest width near midlength; with distinct marginal line basally; 10 complete striae of moderate puncture size; scutellary striole extending 1⁄5 elytral length, with 7–10 punctures; punctures of elytral striae 2× larger than pronotal punctures, striae weakly impressed; intervals of striae with fine, indistinct shallow punctures, 1⁄5 size of strial punctures; all punctures of elytra bearing a single very 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 ½ metaventrite length. Legs narrow, relatively similar in length and shape. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora narrowly oblong, compressed laterally; tibiae shorter than femora, weakly 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 ventrite 5 with distinct denticle at apex. Male genitalia (n = 2) similar to all others in the genus with widened tegmen, elongate cylindrical median lobe, and long coiled flagellum ( Fig. 12G–I ).

Female. Similar to male. Female differs with narrow protibia and protarsus, male have weakly dilated protibial and basal protarsomeres. Female lacking apical denticle on terminal abdominal ventrite. Genital tube shortened, length past abdominal segment VIII = 1.5× width (n = 2); gonostylus set apically on gonocoxite, gonostylus length = 4–5× width ( Fig. 12J ). Spermatheca base rounded on dorsal side, angulate on ventral side, basal third broadest at basal end, narrowing until unsclerotized arc at central third, apical third narrowing and curving gradually to a rounded apex ( Fig. 12K ).

Type specimens. The type specimen(s) of Pharaxonotha kirschii were collected in Silesia on plant materials from Mexico (Reitter 1985; Pakaluk 1988), present location of the type (s) is unknown (not examined). The types of Thallisella condradti were collected in Vera Paz, Guatemala ( Gorham 1898), present location of type material is the NHMUK (examined).

Distribution. As considered here, Paraxonotha kirschii is not associated with cycads and occurs naturally in the wild from the southern US to Panama.

Material examined (total 641). Data presented below were collected over many years of study and specimens returned to the stated repository. Considering the newly recognized cycad species in the kirschii group, most of the following materials need to have their identity confirmed. The only materials confirmed at present are those in the FSCA where all materials not associated with cycads were P. kirschii . Data for materials here considered to be P. kirschii include: BELIZE: Cayo, 0.5 mi. W. Augustine, 30-VII-1976, T. McCarthy, Berlese litter broad-leaf seasonal forest, FM(HD)#76-207 (1 FMNH); Orange Walk, Rio Bravo Cons. Area, vic. LaMilpa Archeol. Site, Site#9, 10–16-IV-1995, P. Kovarik, rotting mushroom baited pitfall (1 FSCA); same data except, Site#12, 11–18- VI-1996 (1 FSCA). COSTA RICA: Cartago: Turrialba, 16–20-V-1979, J.M. & B.A. Campbell (1 CNCI); Guanacaste: 3 km N. Cañas, Hac. La Pacifica, 5–6-VI-1979, H. & A. Howden (2 CNCI); same data except, 9–11- VIII-1987 (1 CMNC); 4 km NW Cañas, La Pacifica, 5-VI-1979, JM & BA Campbell (4 CNCI); Comelco, Palo Verde, OTS, 9 km W. Bagaces, 10°32′N, 85°18′W, 9-IV-1972, J. Wagner, J. Kethley, Berlese, 4 liters conc. organic debris on rocks below falls, drip zone, FM(HD)#73-388, 73CRIV-9a (1 FMNH); Liberia, 20-V-1993, J. & A. Ashe, #047 ex rotting mangos (2 SEMC); Liberia, ca. 32.2 km. N., 23-V-1993, J. & A. Ashe, #046 ex fruit fall litter (40 SEMC, 6 FSCA); Maritza Biol. Sta., 22-V-1993, J. & A. Ashe, #036 ex flight intercept (4 SEMC); same data except, 22-V-1993, #040 ex bracket mushroom (1 SEMC); same data except, 22-V-1993, #039 ex treefall litter (1 SEMC); same data except, 22-V-1993, #038 ex flower fall litter (3 SEMC); Rincon de la Vieja N.P., Las Pailas, 4-IX-1998, C. W. & L. O’Brien (1 FSCA); La Pacifica: Rio Corbili, V– VI-1992, B. Ratcliffe, M. Jameson, C. Dwyer, ex sifting (3 SEMC); Puntarenas: Monteverde, 25-V-1994, J. Ashe, R. Brooks, R. Leschen, Costa Rican Expedition #480 ex black light (1 SEMC); Monteverde Areas, 6-VI-1973, Erwin & Hevel Central American Expedition, 1973 (1 USNM); Monteverde, Pension Quetzal, 14–17-VI-1987, B&B Valentine (1 FSCA); Reserva de Monteverde, 1–4- VI-1979, JM & BA Campbell (1 CNCI); San José: La Caja, III-1929, H. Schmidt leg; F. Nevermann, auf Sandbank (4 USNM); same data except, II-1932 (2 USNM); San Pedro de Montes de Oca, 12-11-1932, C. H. Ballou, C.R.888, reared in raisons (2 USNM); same data except, Rec. Jan 1947, Luis Salas-2, 47-982, corn & beans (14 USNM). EL SALVADOR: Ahuachapan: Parque Nacional el Imposible, Centro de Interpretacion Mixtepec, 13°49′39″N, 89°56′51″ W, V-14-2000, Smith, Ocampo, Cave, Cordero, m. v. light (1 UNSM, 1 MTEC); San Marcos: Los Planes, 25-VI-1959, P. A. Berry (1 USNM); San Salvador, 29-V-5-VI-1958, C. L. Cartwright, at light (1 USNM); Santa Anna: 6.0 km. W. Hwy CA1 above Lago de Coatepeque, 1-VI-1973, Erwin & Hevel Central American Expedition, 1973 (2 USNM). GUATEMALA: [unknown locality], Sallé coll., 1911-403 (1 NHMUK); Alta Vera Paz: Cobán, Condradt, Thallisella condradti Gorham , [red ringed circle] TYPE, B.C.A, Col., VII. (1 NHMUK); San Juan, Champion, 1911-403, Tr. Ent. Soc. L., 1913, det. Champion, Pharaxonotha kirshi R. (see Champion 1913; 1 NHMUK); Senahu, Champion, 1911-403 (1 NHMUK); 22-4, Schwarz & Barber, Cacao Trece Aguas (1 USNM); 17-4, Schwarz & Barber, Cacao Trece Aguas (1 USNM); Baja Verapaz: San Jerónimo, Champion, 1911-403, B.C.A., Col., II(1). Pharaxonotha kirshi Reitt. (1 NHMUK); Guatemala: Capetillo, Champion, 1911-403 (1 NHMUK); Catarina Pinula, Puerta Parada, N145572, W90.4653, 23-II-1-III-2008, J. Schuster (1 FSCA); Jalpa: Mataquescuintla, Finca Concepcion, 2-VII-1986, J. M. Campbell, beating roadside vegetation (1 CNCI); Quiché: Panimá, Champion, 1911-403 (1 NHMUK); Suchitepequez: Finca Chitalon Mazatenango, 8-III-1965, J. M. Campbell (1 CNCI); Finca San Rafael Olimpo, Cuyotenango, 22-I-1966, J. M. Campbell (1 CNCI); same data except, 20-I-1966, ex fungal gardens of Atta nest, 3 ft. below ground level (1 CNCI). HONDURAS: Francisco Morazán: Monte Uycau, 24-V-2993, F. E. Skillman JR, UV & MV light (1 FSCA); Tegucigalpa, El Loarque, 30-VII-1968, B. K. Dozier (2 FSCA); Zamorano, XI-XIII-1970, G. F. Freytag (1 MTEC); Zamorano, 13-X-1993, R. Turnbow (5 RHTC); Zamorano, 24-V-1993, R. Turnbow (1 RHTC); Zamorano, 6-VI-1993, R. Turnbow (1 RHTC); Zamorano, 14°N 87°W, 6-VI-1994, J. Ashe, R. Brooks, #014 ex rotting breadfruit (10 SEMC, 2 FSCA); same data except, 7-VI-1994, J. Ashe, R. Brooks, #028 ex breadfruit fall (1 SEMC); same data except, 7-VI-1994, #056 ex rotting vegetation (1 SEMC); same data except, 9-VI-1994, #055 ex rotting mangos (1 SEMC); same data except, 12-VI-1994, #091 ex rotting flowers (1 SEMC); same data except, 12-VI-1994, #093 ex leaf litter/rotting fruit (1 SEMC); same data except, 12-VI-1994, #094 ex leaf litter along stream (1 SEMC); same data except, 30-VI-1994, #258 ex rotting breadfruit (1 SEMC); Zamorano, along Rio Yeguare, 6-VI-1996, F. G. Andrews & A. J. Gilbert (58 CSCA). MEXICO: Chiapas: 12 mi. E. Santa Isabel at Rio Zoyoltenco, 9-VI-1991, J. Ashe, #81 ex fig-fall and litter (27 SEMC, 2 FSCA); 2 mi. S. Simojovel, 9-VI-1969, J. M. Campbell (1 CNCI); 22 km. N. Ocozocoautla, 1-VII-1969, Campbell & Bright (1 CNCI); same data except, 2-VII-1969 (1 CNCI); 76 km. N. Ocosingo, 1-X-1986, R. Turnbow (1 RHTC); 8 km. S. Chicoasen, 1-VI-1991, J. Ashe, #42, ex leaf litter, sifted (4 SEMC); Barriozabal, VII-1966, ex maiz (4 CSCA); El Chorreadero, 26-VI-1990, R. Turnbow, mv + bl (1 RHTC); Tapachula, 28-II-1990, K. Hibbard, light trap (1 FSCA); Colima: Volcán de Colima, L. Conrad (3 USNM); Volcán de Colima, Conradt (1 USNM); Distrito Federal: Mexico City, Feb-76, F. L. Walters, ex irradiated maize (6 CNCI); Guanajuato: Tupátaro, Höge, B.C.A., Col., II(1). Pharaxonotha kirshi Reitt. (3 NHMUK); Sallé coll., 1911-403 (1 NHMUK); Hidalgo: Jacala, 31-VIII-1960, Howden, at light (1 CNCI); Jalapa: Flohr, 1911-403 (1 NHMUK); Jalisco: 20 mi. SW. Autlan, 13-VII-1982, F. G. Andrews, blacklight (39 CSCA); Nuevo Leon: 10 km. N. Linares, 22-III-1991, R. Brooks, R. Leschen, #57, ex mercury vapor light (3 SEMC); 17 mi. W. Linares, 11-V-1994, J. E. Wappes, 2 FSCA); 29 km. W. Linares, S. Rosa Can., 3-VI-1983, S. & J. Peck, ravine oak forest litter Berlese (1 CNCI); 31.5 km. SW. Linares, 24-III-1991, R. Brooks, R. Leschen, #64, under leaves (1 SEMC); 32 km. SW. Linares, 17-24-III-1991, R. Brooks, R. Leschen, #44 ex flight intercept (10 SEMC); 5 mi. S. Monterrey, 21-VII-1963, H. F. Howden (1 CNCI); same data except, 6-VI-1963 (1 CNCI); same data except, 31-VI-1963 (1 CNCI); same data except, 20-VI-1963 (1 CNCI); Monterrey, 27-IV-1969, H. F. Howden (1 CNCI); Monterrey, Chipinque Mesa, 21-25-VI-1969, S. & J. Peck, forest (1 CNCI); Oaxaca: Tlaxiaco, 3-XII-1902, A. L. Herrara, in corn (7 USNM); Mitla, 27-VI-1900, C. C. Dean (1 USNM); Chittenden coll [likely an interception] (4 USNM); Querétaro: Arroyo Seco, San José de la Flores, 21°22′52, N 99°30′12, W, 6-VII-2003, J. A. Peréz de la Rosa y G. Vargas, ex Dioon edule (cono femenino) (1 CZUG); Mpio.Jalpan, RT-190, 4.9 km. NW. Zoyapica, 21°20′58″N, 99°18′38″W, 11-XII-2013, Kovarik, Skelley, Jones, ex oak leaf litter Berlese (4 FSCA, 2 UAQM); Tamaulipas: Bocatoma w.s., 7 km. SSE Gomez Farias, 15-X-1985, R. Turnbow (1 RHTC); Cd. Mante, 27-VII-1960, H. Howden, light (1 CNCI); Guemes, 15 km. N. Ciudad Victoria, 6-VI-1961, Univ. Kans. Mex. Exp. (14 SEMC); 12 miles south of El Mante, 17-VI- 1941, H. Dybas (1 FMNH); Veracruz: 2.5 km. S. Jalapa, 28-V-1991, J. Ashe, #26 ex fungusy log (1 SEMC); Córdoba, 21-VII-1941, H. Dybas, at light (1 FMNH); Córdoba, Höge, B.C.A., Col., II(1). Pharaxonotha kirshi Reitter (2 NHMUK); Córdoba, Sallé coll. (3 NHMUK); Rd 185, 2 mi. S. Acayucan, 10-VII-1962, Campbell & Hill (2 CNCI); Tuxtla, Sallé coll., 1911-403 (1 NHMUK). NICARAGUA: Granada: Reserva Silv. Priv. Domitila, 11°42.50′N, 85°57.20′W, 6-9-VI-2002, R. Brooks, Z. Falin, S. Chatizimanolis, ex flight intercept trap NIC1BFC02_255 (1 CMNC); Reserva Silv. Priv. Domitila, 11°2.322′N, 85°6.884′W, 31-V–6-VI-2005, Fred. G. Andrews, mercury vapor light (3 CSCA); Reserva Silv. Priv. Domitila, 11.70830°, −85.95330°, 31-V-2011, C. W. Shin, NIC-CWS-2011 001 (43 SEMC, 8 FSCA); same data except, 1-VI-2011, at light, NIC-CWS-2011 007 (12 SEMC); same data except, 2-VI-2011, at light, NIC-CWS-2011 016 (22 SEMC); same data except, 4-6-VI-2011, Malaise trap, NIC-CWS-2011 Mal03 (1 SEMC); same data except, 6-VI-2011, Mercury vapor light on hill, NIC- CWS-2011 021 (13 SEMC); same data except, 8-10-VI-2011, flight intercept trap near stream, NIC-CWS-2011 FIT05 (2 SEMC); same data except, 9-VI-2011, Mercury vapor light, NIC-CWS-2011 066 (87 SEMC); same data except, 13-VI-2011, Mercury vapor light, NIC-CWS-2011 090 (1 SEMC); same data except, 14-VI-2011, Mercury vapor light, NIC-CWS-2011 092 (1 SEMC); Managua: Managua, 4-VI-2018, R. Turnbow (1 RHTC); Rio San Juan: 60 km. SE. San Carlos, Refugio Bartola, 10°58.40′N, 84°20.30′W, 26-V-2002, R. Brooks, Z. Falin, S. Chatzimanolis, ex ginger fruits, NIC 1BFC02 077 ( SEMC). PANAMA: Canal Zone, 12-V-1952, F. S. Blanton, at light (1 USNM); Chiriquí: Chiriquicito, 18-V-1996, R. Turnbow, mv + bl (1 RHTC); Coclé: 5.6 km. N. Pan Amer. Hwy, El Copé Rd, 08°37′N, 80°35′W, 7-VI-1995, J. Ashe, R. Brooks, #141 ex slash (2 SEMC); Panama: 11–15 km. N. El Llano, 13-V-1991, R. Turnbow (1 RHTC); Fort Kobbe, 27-V-1965 (1 USNM); same locality, 12-V-1985, A. J. Gilbert, P. H. Sullivan, and F. T. Hovore (1 FSCA); same locality, 23-V-1991, R. Turnbow, mv + bl (1 RHTC). USA: Louisiana: Vernon Parish, nr. Rosepine, Castle Farm Site 1, N30.89628, W93.33945, 30-VI-2005, S. T. Dash, W. Lomak, L. Cannizzaro, open pasture w/white oak nearby on hill, pitfall tran. PF1-8 (2 FSCA); Texas: [?County], San Antonio, 1-12, Hubbard & Schwarz (1 USNM); Anderson Co., Tennessee Colony, 4–5-III-1967, A. Blanchard (1 USNM); Bastrop Co., Bastrop State Park, 14-IV-1959, Baker & Howden (2 CNCI); Bee Co., Beeville, 5-11-53, Univ. Kans. Mex. Exp. (1 SEMC); Bexar Co., 3 mi. N. Leon Springs, Scenic Oaks, 29°41′21″N, 98°39′40″W, 31-III- 2012, J. E. Wappes, UV light trap (1 FSCA); same data except, 1-IV-2012 (1 FSCA); same data except, 30-IV-2012 (3 FSCA); same data except, 21–25-IV-2015 (12 FSCA); Bexar Co., Leon Valley, 14-VI-1971, G. H. Nelson, blacklight (2 FSCA); same data except, 3-VII-1971 (1 FSCA); Caldwell Co., 4.5 mi. E. McMahan, 1-VI-1998, Wappes & Turnbow, MV/UL (1 FSCA); Comal Co., vic. Bulverde, 31-V-1998, JE. Wappes (1 FSCA); Cottle Co., Paducah, 9-VII-1967, A. & M. E. Blanchard (1 USNM); Guadalupe Co., 10 mi. SE. Seguin, 1187 Kolman Rd., 29°29′N, 97°51′W, 17-21-V-2014, J. E. Wappes, MV/UV (1 FSCA); same data except, 23-IV-2015 (1 FSCA); Kerr Co., Kerrville, 5-VI-1956, H. & A. Howden, light (2 CMNC); same data except, 22-VII-1956 (2 CMNC); Kerr Co., Kerrville, 1-IV-1959, Becker & Howden, in malt trap (12 CNCI); same data except, 4-IV-1959 (1 CNCI); same data except, 5-IV-1959 (1 CNCI); same data except, 6-IV-1959 (2 CNCI); same data except, 10-IV-1959 (2 CNCI); same data except, 13-IV-1959 (6 CNCI); Kimble Co., Junction, 29–30-1967, A. & M. E. Blanchard (1 USNM).

Intercepted materials (total 33). These data represent specimens collected in commercial transport into Canada and the USA. They are deposited in the California ( CSCA), Canada ( CNCI), and United States ( USNM) institutions with identification services for their official agricultural inspectors. There are no associated data that indicates where in US or Canada these were intercepted. Label data with host information include specimens originating: From Guatemala: 19-VI-1935, H. Y. Gouldman, E.Q.A31625, with corn (1 USNM). From Mexico: VI-1925, #277, on pke of cotton seed meal (1 USNM) ; VIII-1929, S.F.2236, corn (1- CSCA) ; 6-IX-1933, L.J. Ray , #1401, in cotton rabbits (1- USNM) ; 17-VII-1934, H. Y. Gouldman , B.P.Q. A29060 , on tree seeds (3 USNM) ; 22-VI-1935, G. F. Callingham , #1958, on pineapple (1 USNM) ; 7-VII-1935, L. R. Dorland Thayer , 209, on sugar cane (1 USNM) ; 30-III-1936, 2881, on beans (1 USNM) ; 23-IX-1936, H. A. Carey , #7078, 36-32095, on cacti (6 USNM) ; 2-X-1936, B. C. House , #7097, 36-32096, on cactus plants (5 USNM) ; 8-X-1936, #7168, on cactus (1 USNM) ; 8-IX-1942, No. 29673, orchids (1 USNM) ; 12-VII-1944, 44-19882, on lilies (1 USNM) ; 7-XII-1950, Zuck colr. #72490, lot 52-13872, with dry banana fruit (2- USNM) ; 22-V-1952, H. Y. Gouldman , X28238 , lot-52-6282, Cucurbita moschata seed (1 USNM) ; 4-XI-1955, D. Brock, Cal. Dept. Agr. No. 55K55, on corn (2 CSCA) ; 1957, E. de las Casas , en granos almacenados (2 USNM) ; 8-VIII-1966, ex corn (4 CNCI). From southern USA: XII-1943, ex peanuts from El Paso, Texas (1 CNCI) .

Remarks. Based on the volume of data for P. kirschii listed above, it appears to be a more free-living species, occurring more commonly in leaf litter in the wild in many areas where there are no native or ornamental cycads. All specimens intercepted on plant products as a pest were associated with various grains or tubers. Even with the recognition of cycad associated species in the kirschii group, it is possible that cryptic species reside within the materials here considered P. kirschii . We prefer to leave their analysis for future work when molecular grade materials are available for study.

Pharaxonotha kirschii was described from specimens collected in eastern Europe from Mexican plant products. Since then, it has been intercepted in many countries and mentioned many times in regional lists, regional reports, taxonomic catalogues, phylogenetic studies, keys to taxa, natural history accounts, etc., and now a growing number of websites. Many references cite the genus and species names with different spellings (see Leschen and Węgrzynowicz 1998). An effort was made to find every reference listing the species which might present additional information. Most only list the name or repeat previous information and are too numerous to present here in a comprehensive accounting. In the following paragraphs, we review some of the important literature.

In the original description, Reitter (1875) stated P. kirschii was collected in Opava [Troppau], Czechoslovakia, in a plant product [depending on the translation it could be drugs, grain, or spelt] from Mexico. That was followed by the accounts of Riley (1894) and Chittenden (1895) discussing damage to grains at the World’s Fair, Columbian Exposition in 1893, Chicago, Illinois, USA. Chittenden (1911), who possessed a captive colony, described and illustrated the life stages and made observations on the life history of P. kirschii . Larvae were illustrated again by Böving and Craighead (1931), and are only mentioned in other references (e.g., Roberts 1939; Sen Gupta and Crowson 1967, 1971; Lawrence 1991). With one exception, all literature with life history information simply repeats Chittenden (1911), this includes Hinton (1945) and Booth et al. (1990). More recent original work on their life history was published by Konoike et al. (1987).

The description of P. kirschii was soon followed by the description of Pharaxonotha zamiae Blake, 1928 [now a synonym of P. floridana ( Casey, 1890) by Kingsolver 1973], that was reared from the “flowers of Zamia floridana ”. After 60 years with no new species described in Pharaxonotha, Pakaluk (1988) offers a short redescription of P. kirschii while describing two new cycad associated species in the genus. With several species in the genus being associated with cycads, there has been a misconception that P. kirschii could also be a cycad associate in the wild. We now know this is not the case.

As the “Mexican grain beetle”, P. kirschii has been discussed in references including Chittenden (1895, 1911), Hinton (1945), Anderson (1987), Kingsolver (1987), Booth et al. (1990), Lawrence (1991) and many others. They report it feeding on stored plant products such as corn, corn meal, wheat, flour, beans, and edible tubers. Based on label data in this study, we can add that it has also been associated with imported cactus, orchids, fungus logs, lilies, pineapples, cotton, cotton seed meal, sugar cane, peanuts, squash ( Cucurbita moschata Duchesne ex Poir ) seeds, and tree seeds. Wild collections indicate associations with fruit, flower and treefall litter, leaf litter, various rotting fruits, rotting mushrooms, bracket fungus, fungusy log, leaf cutter ant fungal gardens, and general organic debris.

With the recognition of the cycad associated species in the kirschii group, we know of only one specimen of true P. kirschii associated with a cycad. This specimen was collected with the three specimens of P. sclerotiza on D. angustifolium in Querétaro. Our concept of P. kirschii is based on the type of P. condradti and specimens collected with stored products or collected in the wild without any cycad association. It is also possible that these non-cycad associated materials harbor cryptic species.

Pharaxonotha kirschii superficially resembles some species of Cycadophila Xu et al. in Asia, a genus that also has members associated with cycad pollination and others that are not ( Xu et al. 2015; Skelley et al. 2017). We speculate that there is convergence in body characters for Pharaxonotha species that occur in early vs. late cone stages of development or are free living. This would allow for a stratification of niches. Species of Cycadophila associated with later cone development or not associated with cycads are similar externally to the kirschii group. While those occurring on early cone development and possibly acting as primary pollinators are superficially more similar to other Pharaxanotha with similar habits. Diet shifts within in the Erotylidae were studied by Leschen and Buckley (2007), who briefly mention Pharaxonotha as cycad feeders.

Figures

Figure 1. Pharaxonotha heads, lateral view. A) P. kirschii, Mpio. Jalpan, Querétaro, Mexico. B) P. bicolor, paratype, San Andres Pápalo, Oaxaca, Mexico. C) P. occidentalis, paratype, El Campanario, Chiapas, Mexico. D) P. vovidesi, paratype, Mt. Oscuro, Veracruz, Mexico.

Figure 2. Pharaxonotha body parts. A–C) Dorsal view of head. A) P. kirschii, Mpio. Jalpan, Querétaro, Mexico. Arrow points to temple. B) P. occidentalis, paratype, El Campanario, Chiapas, Mexico. C) P. vovidesi, paratype, Mt. Oscuro, Veracruz, Mexico.D) P. kirschii, abdominal ventrites, Mpio. Jalpan, Querétaro, Mexico. Arrow points to tooth on ventrite 5. E–F) Antennal clubs. E) P. kirschii, Santa Isabel, Chiapas, Mexico.F) P. sclerotiza, holotype. G) Spermatheca of Pharaxonotha fawcettae from Jalpan de Serra, Querétaro, Mexico indicating the diagnostic structures and orientations used in descriptions.

Figure 12. Pharaxonotha kirschii from Bexar County, Texas, USA. A–C) Male dorsal, lateral and ventral habitus. D–F) Male head, dorsal, lateral and ventral views. G) Tegmen attached to median lobe and genital capsule, ventral view. H) Tegmen attached to median lobe, lateral view. I) Parameres of tegmen, ventral view top and lateral view bottom. J) Spermatheca and retracted ovipositor, ventral view. K) Spermatheca (partially inflated).