Verodes Casey, 1907

Foley, Ian A. & Ivie, Michael A., 2008, A phylogenetic analysis of the tribe Zopherini with a review of the species and generic classification (Coleoptera: Zopheridae), Zootaxa 1928 (1), pp. 1-72 : 49-53

Genus Verodes Casey NEW SENSE

Figs. 31 , 34 , 55–68

Homaloderes Solier, 1834: 31 (nomen nudum).

Nosoderma Solier, 1841: 31 , not Guérin-Méneville, 1838: 280 [homonomy discussed by Foley and Ivie 2007] LeConte, 1851: 130. LeConte, 1853: 235. LeConte, 1859 a: 77. LeConte, 1859 b: 655. LeConte, 1862: 216. Kirsch, 1866: 189. Pascoe, 1866: 487. Champion, 1884: 44. Fairmaire, 1894b: 656. Casey, 1907a: 44. Casey, 1907b: 470. Gebien, 1936: 669. Blackwelder, 1945: 514. Triplehorn, 1972: 1. Doyen and Lawrence, 1979: 341. Lawrence, 1991: 518. Lawrence and Newton, 1995: 891. Ṡ lipi ń ski and Lawrence, 1999: 13. García-París et al., 2000: 473. García-París et al., 2001: 144. Ivie, 2002: 460. Garrido, 2004: 56. García-París et al., 2006: 215-230. (Type species Nosoderma morbillosum Solier by subsequent designation of Gebien, 1936).

Verodes Casey, 1907b: 470 . Gebien, 1936: 670. Neave, 1939: 635. Blackwelder, 1945: 515. Foley and Ivie, 2007: 69 . (Type species Nosoderma aequalis Champion by original designation, Synonymy by Doyen and Lawrence, 1979: 342.).

Noserodes Casey, 1907b: 470 Gebien, 1936: 670 . Neave, 1939: 346. Blackwelder, 1945: 515. (Type species Noserodes squalidus Casey, 1907b: 480 by original designation). (Synonymy by Doyen and Lawrence, 1979).

Diagnosis: The Verodes clade as recognized here is supported by the presence of a distinct membranous pros-

theca on the mandible ( Fig. 34 ), ventrally inserted labial palps ( Fig. 56 ), lack of an antennal cavity on the

hypomeron (often a depression, or a small notch), and the presence of tubercles on the ventral surface (also in

Phloeodes ).

Description: See Ṡ lipi ń ski and Lawrence (1999). The previously published description of the genus

Nosoderma is accurate as here classified based on the removal of two species to Sesaspis and one to Phloe-

odes.

Notes: The genus Verodes is the second largest in the tribe behind Zopherus , and prior to this study contained 23 valid named species ( sensu Foley and Ivie 2007 ). Morphological analysis of this group suggests that it is not monophyletic. The species group centered on Verodes denticulata forms a distinct group and is here recognized as the reconstituted Casey genus Sesaspis NEW SENSE. When Sesaspis is recognized as a genus, Verodes , Phloeodes , and Sesaspis all form monophyletic genera with the only remaining problem being the placement of Verodes venustus . This species does not fit the definition of either Verodes or Sesaspis . It is excluded from Verodes because it lacks a membranous mandibular prostheca, and from Sesaspis because the ventral surface is densely tuberculate and it lacks an arcuate groove on the hypomeron. Based on this evidence, Verodes venustus is moved to Phloeodes NEW COMBINATION, and becomes the only member of that genus to occur outside the California Floristic Province.

The genus Verodes as here defined contains 11 species in two separate morphological clades. A group of seven species occurs generally from Oaxaca, Mexico, north possibly into the United States (there are unconfirmed localities of Verodes exsculptus in Texas with simply state data [NMNH]), and another group containing four species that generally occur from Oaxaca, Mexico, south into Central America.

The species-group occurring northwards from Oaxaca is generally restricted to high-elevation cloud forests and contains the two most widespread and morphologically diverse species of Verodes , V. inaequalis and V. exsculptus , and five that are quite rare. Verodes championi , the only species in the genus described by Casey that appears valid, is similar in appearance to V. inaequalis , but has a unique elytral sculpture that approaches the pattern of V. exsculptus . Verodes carinatus is most similar to V. exsculptus but can be distinguished by the stout body form and differing elytral sculpture. Verodes insignis is known only from the state of Oaxaca, and the elytral sculpture of this species, consisting of ridges that are hooked at the apex, is unique in Verodes . Verodes scabrosus and V. asperatus appear to be sister-species that are isolated to high elevation cloud forests.

The lineage occurring in more southern locales of Mexico and into Central America contains the species that Casey placed into the genera Noserodes ( V. zunilensis , V. guatemalensis , and V. sparsus ) and Verodes (sensu Casey 1907b) ( V. aequalis ). The three species previously placed in Noserodes are closely allied, and V. zunilensis and V. guatemalensis are probably sister-species. The species formerly placed in its own genus by Casey, V. aequalis , is distinct among species of Verodes in having the elytral sculpture completely reduced, even more so than Sesaspis doyeni and S. ashei , and lacking nodules on all femora of the male, but these characters do not warrant generic distinction based on the current analysis.

Most species of Verodes are highly variable in morphology, especially in the intensity of the nodules and ridges that form the elytral and pronotal sculpture. The key below uses this sculpture extensively which makes the identification of a single specimen more difficult than several representatives of a species.

Distribution: Many specimens of this genus were collected in the days of the Biologia Centralia-Americana during the 19 th century and therefore often lack accurate locality information. The majority of specimens have simply “ Mexico ” or a few localities common to the above mentioned publication such as “Jalapa”, “Las Vigas”, “ Guerrero ”, and others, but a general distribution pattern for the genus does appear to form. The majority of the diversity in the genus occurs around Oaxaca, Mexico, with radiations north though the Sierra Madre Occidental and south along the western coastal ranges of Central America. Interestingly, the genus Sesaspis (although the genera do co-occur at the same localities in Mexico) appears to occur in the other half of Mexico radiating through the Sierra Madre Oriental, and occurring in the lowland tropical forests of the eastern edge of Central America. Some of these distributional patterns were recognized by García-París et al. (2001, 2006) but they did not recognize Sesaspis as an independent lineage.

Figures

FIGURES 30–31. Ventral surface of the head. 30. Sesaspis sylvatica; 31. Verodes aequalis. Scale bars = 1mm.

FIGURES 32–34. Dorsal view of left mandibles. 32. Phloeodes diabolicus; 33. Sesaspis sylvatica; 34. Verodes aequalis. Scale bar = 1mm.

FIGURES 54–56. 54–55. Hypomera. 54. Sesaspis sylvatica; 55. Verodes exsculptus. 56. Ventral surface of head and pronotum of Verodes inaequalis. Scale bars = 1mm.

FIGURES 57–60. Dorsal habiti. 57. Verodes inaequalis, Mexico, Puebla.; 58. V. exsculptus, Mexico, Oaxaca; 59. V. asperatus, Mexico, Hidalgo; 60. V. scabrosus, Mexico. Scale bars = 5mm.

FIGURES 61–64. Dorsal habiti. 61. Verodes interruptus, Mexico, Oaxaca; 62. V. sparsus, Guatemala; 63. V. zunilensis, Mexico, Chiapas; 64. V. guatemalensis, Mexico, Chiapas. Scale bars = 5mm.

FIGURES 65–68. Dorsal habiti. 65. Verodes impressus, Mexico, Veracruz; 66. V. carinatus, Mexico, Veracruz; 67. V. championi, Mexico, Oaxaca;68. V. aequalis, Mexico, Oaxaca. Scale bar = 5 mm.