Pharaxonotha vovidesi Skelley, Tang and Pérez-Farrera, 2022

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 : 14-17

Pharaxonotha vovidesi Skelley, Tang and Pérez-Farrera , new species

Figures 1D , 2C , 5A–K

Diagnosis. Pharaxonotha vovidesi is distinguished from other species in the genus by the pronotum and elytra with long, distinct setae; head flattened anteriorly and convex between eyes, head with transverse occipital line short, indistinct laterally, sexual dimorphisms indistinct, and pronotum with mean PL/PW ranges from 0.71–0.74 (multiple specimens required). In females the spermatheca is short and compact, ratio of spermatheca length to width at basal third <3.5, and base abruptly swollen, swelling often quadrate-shaped. It is known to inhabit D. edule south of the Mexican Transvolcanic Belt in Veracruz, Mexico.

Description. Length 2.48–3.80 mm, width 0.96–1.47 mm. General body color entirely pale yellowish brown ( Fig. 5A–C ); dorsal surface punctate, strongly alutaceous, dulled, long procumbent hairs associated with punctation on pronotum and elytra; ventrally alutaceous, dull, covered with scattered long procumbent setae.

Head. Not broad, width = 0.67–0.77× pronotal width ( Fig. 5D–F ); in lateral view flattened anteriorly, convex between eyes; in dorsal view conical, gradually narrowed anteriorly, moderately punctured, average distance between closest punctures 1–2× width of puncture; head width 0.57–0.81 mm; dorsal interocular distance 0.40– 052 mm, head width/dorsal interocular distance ratio 1.48–1.65, ventral interocular distance 0.30–0.39 mm, head width/ventral interocular distance ratio 1.94–2.29. Eye with large black facets, about 2× diameter of head punctures. Antennal length slightly longer than pronotal width, 1.5× head width; antennomere I (scape) fairly large, slightly elongate; antennomere II slightly shorter than III; IV 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] short laterally in most, most with line stopping about half distance between stridulatory file and eye, not curving anteriorly. Mentum and submentum moderately punctured, 1/3–½ diameter of facet, distance between nearest punctures approximately 1–2× own diameter, each puncture with a long seta. Gular area smooth, without punctation or setae, border with submentum marked by change in punctation and with a shallow transverse depression.

Thorax. With pronotum rectangular in dorsal view, length/width ratio 0.67–0.77; with distinct marginal carina laterally and basally, anteriorly lacking marginal carina medially; surface convex; anterior angles narrowly rounded in dorsal and lateral view, projecting forward; posterior angles obtuse, with small denticle at angle; lateral margin 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 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 convex apically, weakly expanded and truncate at apex. Hypomeron smooth, with indistinct seta bearing punctures. Scutellar shield distinctly transverse, posterior margin weakly roundly pentagonal. Elytra in dorsal view elongate-oval, convex; length/width 1.69–1.88, greatest width near midlength; with distinct marginal line basally; 10 complete striae of moderate puncture size; scutellary striole extending ¼ elytral length, with 10–15 punctures; punctures of elytral striae as large as pronotal punctures, striae not impressed; intervals of striae with fine, distinct punctures, ½ size of strial punctures; all punctures of elytra bearing a single prominent long seta, extending past next puncture in series. Mesoventrite with strong punctation, distance between nearest punctures approximately equal to diameter of punctures, puncture depth moderate. Metaventrite weakly glossy, with moderate lateral punctation separated by 2–3× own diameter; medial surface finely punctured, separated by 3–4× own diameter; entire surface convex, metathoracic discrimen extending approximately ¾ metaventrite length. Legs narrow, relatively similar in length and shape. Procoxa oval; mesocoxa globular; metacoxa transversely elongate-oval; trochanters obliquely truncate apically; femora narrowly robust, moderately compressed laterally; tibiae shorter than femora, narrowly dilated to obliquely truncate apices; protibia with apical lateral tooth small, distinct, with apical fringe of long stout spinules on concave ventral apical margin slight gap present between last spinule and lateral tooth; meso- and metatibia with apical fringe of long spinules on anterior and posterior margins.

Abdomen. Ventrite apical margin bearing long setae; all ventrites bearing moderate, shallow evenly distributed 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 4× diameter of puncture; I–IV each with 2 or more median pairs of longer, semi-erect sensory hairs, they are obscured in other setation. Male genitalia (n = 6): not distinctly dorsoventrally flattened; tegmen in dorsal view with basal piece ring-like; parameres in dorsal view narrowed at center, apical half slightly asymmetric with slight bulge on outer margin, in lateral view length = 3.8–4.5× width; elongate cylindrical median lobe narrowing apically; long coiled flagellum ( Fig. 5G–I ).

Female. Similar to male, except male femora, tibia and protarsi slightly more robust. Female genitalia (n = 8): gonostylus cylindrical, widest at apex, tapering slightly and gradually to base, set apically on gonocoxite, gonostylus length = 3.3–4.0× greatest width ( Fig. 5J ). Spermatheca C-shaped, slightly asymmetric, widest in basal third, length = 3.0–3.4× greatest width, base abruptly swollen, swelling often quadrate-shaped, basal third with longitudinal wrinkles, apical third weakly annulated ( Fig. 5K ).

Distribution. As currently circumscribed, this species is known to occur near Actopan, Colorado, El Farallón and Monte Oscuro in eastern Veracruz, Mexico.

Material examined. Holotype (by designation) male of Pharaxonotha vovidesi with the following labels: 1) [rectangular; white; printed in black ink] “ MEXICO, Veracruz, Monte Oscuro , ex Dioon edule ♂ cone, 2-XI-2012, A. Vovides; 2) [rectangular; red; printed in black ink] “ HOLOTYPE ♂ Pharaxonotha vovidesi Skelley, Tang and Pérez-Farrera 2022 ”. Deposited in the FSCA.

Paratypes (total 274). Allotype ( FSCA) and 183 paratypes with same data as holotype: MEXICO: Veracruz: Monte Oscuro, Municipio Emiliano Zapata, 20-VIII-1993, A. Vovides, C. Iglesias & P. Aguilar, Dioon edule male cone (9); same locality except, X-2002, A. Vovides, 34A, Dioon edule (8); same locality except, X-2002, A. Vovides, C. Iglesias & P. Aguilar, Dioon edule male cone (73). Paratypes to be deposited in ANIC, CUIC, CZUG, FSCA, IEXA, NHMUK, NZAC, UAQM, UNAM, USNM.

Additional specimens (total 308). Material studied presently considered to be P. vovidesi , but not designated as paratypes: MEXICO: Veracruz: Mun. Actopan, Rancho del Niño, 8-X-2002, A. Vovides, C. Iglesias, & P. Aguilar, Dioon edule (66); Colorado, [GPS omitted], Dioon edule ♂ cone, 17-XI-2014, W. Tang (14); Farallón, [GPS omitted], ex ♂ cone Dioon edule , 16-XI-2012, W. Tang (122). These will be deposited in institutions stated above.

Etymology. The species honors Andrew Vovides for his accomplishments in cycad taxonomy, systematics, ecology and conservation as well as his contributions to the study of cycad pollination.

Remarks. Genetic analysis of the host Dioon populations south of the Mexican Transvolcanic Belt by Gutiérrez- Ortega et al. (2018) suggests long genetic isolation of different populations, despite relatively close geographic proximity. We consider P. vovidesi as a complex in which varieties or possibly subspecies may be recognized in the future, if or when more detailed analyses become available. Pharaxonotha vovidesi has been found together on male Dioon edule cones with species of Allocorynina weevils: Parallocorynus (P.) perezfarrerai Tang and O’Brien and P. (Neocorynus) iglesiasi Tang and O’Brien ( O’Brien and Tang 2015) .

Occidentalis species group

Adult diagnosis. In the occidentalis species group the head is more flattened in lateral view, even between the eyes, the transverse occipital line [vertexal line] is absent (except in P. gigantea ), sexual dimorphisms more pronounced, and the dorsal surface is uniformly yellow-brown to orange-brown, with the pronotum often being slightly darker than the elytra. As in the majority of Pharaxonotha inhabiting Dioon , the dorsal surface of the head, pronotum and elytra have long setae emerging from punctures, with setal length reaching or exceeding the distance to nearest puncture (slightly shorter in P. gigantea ). In Pharaxontha inhabiting other cycad genera, dorsal hairs are shorter, not reaching the nearest puncture.

Remarks. This species group is proposed based on analysis of the 16S rRNA gene of three populations of one species that are part of the early diverging lineages in Tang et al. (2018b; 2020). Materials of P. occidentalis analyzed by Tang et al. (2018b: 8, fig.4; 2020: 18, fig. 3) are presented in their trees labeled as follows: “D0177 Loxicha, D. holmgrenii ”; “D0246, Rancho Limon, D. holmgrenii ”; “D0286 Campanario, D. merolae ”.

Four species are placed within the group based on geographic proximity and selected morphological characters. This group is distributed along the Pacific drainage of western Mexico, in the states of Chiapas, Guererro, Jalisco, and Oaxaca on Dioon holmgrenii , D. merolae , D. stevensonii and D. tomasellii and includes one member in eastern Oaxaca, on D. spinulosum , which is within the Atlantic drainage of Mexico. It is likely that members of this species group also inhabit Dioon in other western states of Mexico, such as Durango, Michoacán, Nayarit, Sinaloa, and Sonora. Dissected adult specimens from all four species of the occidentalis group usually have their guts packed with the distinctive monosulcate pollen grains of cycads, indicating that this is a major food source.

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 5. Pharaxonotha vovidesi from Monte Oscuro, Veracruz, Mexico (A–D, F–I: male holotype). A–C) Dorsal, lateral and ventral habitus. D–F) Head male, dorsal, lateral and ventral views. G) Tegmen attached to median lobe, ventral view top and lateral view bottom. H) Parameres of tegmen, ventral view top and lateral view bottom. I) Male genital capsule, ventral view. J) Spermatheca with retracted ovipositor, ventral view. K) Spermatheca (fully inflated).