Phloeodes diabolicus (LeConte)

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 : 42-45

Phloeodes diabolicus (LeConte)

Figs. 32 , 35–36 , 39, 41 , 44 , 47 , 50–51

Nosoderma diabolicum LeConte, 1851: 130 . LeConte, 1853: 235. LeConte, 1857: 49, pl. 2. LeConte, 1859: 77. Pascoe, 1866: 487. Heyden, 1885: 307. García-París et al., 2006: 229.

Phloeodes diabolicus: Horn, 1870: 273 . Casey, 1907a: 42. Leng, 1920: 223. Essig, 1926: 429. Gebien, 1936: 667. Doyen, 1976: 267–272; fig. 1–11. Powell and Hogue, 1979: 295, 297. Arnett, 1983: 16. White, 1983: 250, fig. 107. Costa et al., 1988: 213. Ṡ lipi ń ski and Lawrence, 1999: 19, figs. 69–81. Ivie, 2002: 458, 461.

Nosoderma diabolica: Casey, 1907b .

Ageonoma diabolica Pascoe, 1866: 487 .

Nosoderma pustulosum LeConte, 1859: 77 . García-París et al., 2006: 230.

Phloeodes pustulosus: Horn, 1870: 273 . Casey, 1907a: 43. Leng, 1920: 223. Essig, 1926: 428. Gebien, 1936: 668. Doyen, 1980: 3. Arnett, 1983: 16. White, 1983: 250. Hogue, 1993: 296, 298. Ivie, 2002: 461. NEW SYNONYMY.

Phloeodes latipennis Casey, 1907a: 43 . Casey, 1907b: 472. Leng, 1920: 223. Gebien, 1936: 668. Arnett, 1983: 16. (Synonymy by Gebien 1936)

Phloeodes ovipennis Casey, 1907a: 42 . Casey, 1907b: 472. Leng, 1920: 223. Gebien, 1936: 667. Arnett, 1983: 16. (Synonymy by García-París et al. 2006)

Phloeodes elongatus Casey, 1907a: 42 . Casey, 1907b: 472. Leng, 1920: 223. Gebien, 1936: 668. (Synonymy by García- París et al. 2006).

Phloeodes scaber Casey, 1907a: 43 . Casey, 1907b: 472. Leng, 1920: 223. Gebien, 1936: 668. Arnett, 1983: 16. (Synonymy by García-París et al. 2006).

Phloeodes angustus Casey, 1907a: 43 . Casey, 1907b: 472. Leng, 1920: 223. Gebien, 1936: 668. Arnett, 1983: 16. (Synonymy by García-París et al. 2006).

Phloeodes remotus Casey, 1907b: 472 . Leng, 1920: 223. Gebien, 1936: 668. Arnett, 1983: 16. (Synonymy by García- París et al. 2006).

Diagnosis: The presence of large antennal cavities that are clearly limited posteriorly and completely contain the antennae while in repose ( Figs. 39, 41 ), a few arcuate velvety spots on the elytra, and the reduced pattern of elevated ridges to at most one small nodule at the start of the declivity will distinguish individuals of the species from all other Zopherini with 10-segmented antennae.

Description: See García-París et al. (2006).

Distribution: California Floristic Province as far north as Shasta county, possibly all the way to Umatilla county in Oregon (3 specimens: 2-OR label only [NHMB, FMNH], 1-OR: Umatilla [UGCA], and east possibly to the edge of Arizona (6 specimens: 4-AZ label only [INHS, CASC, MSUC], 1-Yuma [INHS], 1-“Grand Canon”/ VII-1924; Van Dyke [CAS]), and south into California. Specimens have also been recorded from Alaska, Oklahoma, Oregon, Pennsylvania, Texas and Wisconsin though none are from specific localities and can almost certainly be discounted as labeling errors, or hitchhiking individuals.

Recorded distribution: A summary of the 3,878 specimens examined is (for complete label data, see Foley 2006) MEXICO: BAJA CALIFORNIA: Ensenada. UNITED STATES: ARIZONA: Yuma. CALIFOR- NIA: Alameda, Amador, Calaveras, Contra Costa, El Dorado, Fresno, Glenn, Humboldt, Imperial, Kern, Kings, Lake, Los Angeles, Madera, Mariposa, Merced, Mono, Monterey, Napa, Orange, Placer, Riverside, Sacramento, San Benito, San Bernadino, San Diego, San Joaquin, San Jose, San Luis Obispo, Santa Barbara, Santa Clara, Santa Cruz, Shasta, Solano, Stanislaus, Tulare, Tuolumne, Ventura, Yolo. OREGON: Umatilla .

Types: Nosoderma diabolicum : LECTOTYPE here designated: ♂ intact. Hand written Phloeodes diaboli- cus (LeC.)/ Red square Type-4505/ gold circle/ Jan.-Jul MCZ image database/ red rectangle Lectotype; Nosoderma ♂; diabolicum LeConte 1851 ; designated by I. A. Foley 2006.

Nosoderma pustulosum : HOLOTYPE ♀, missing left meso, and right meta tarsomeres in MCZ.. Red square Type 4506/ hand written P. pustulosus : (LeC.): Tejon/ Jan.-Jul. 2005 MCZ Image Database.

Phloeodes latipennis : LECTOTYPE, here designated: ♂, intact in NMNH. Cal.(ifornia)/ Casey; bequest; 1925;/ orange rectangle Type NMNH; 46376/ hand written latipennis; Csy./ Casey determ.; pustulosus-10/ red square Lectotype; Phloeodes ; ♂ latipennis ; Casey 1907; designated I. A. Foley 2006.

Phloeodes ovipennis : LECTOTYPE, here designated: ♀, right pro leg missing last two tarsomeres in NMNH. Cal.(ifornia)/ Casey; bequest; 1925;/ orange rectangle Type NMNH; 46375/ hand written ovipennis Csy. / Casey determ.; elongatus- 4/ red square Lectotype; Phloeodes ; ♀ ovipennis ; Casey 1907; designated I. A. Foley 2006. PARALECTOTYPE, here designated: ♂, intact in NMNH. Cal.(ifornia)/ Casey; bequest; 1925/ orange rectangle ovipennis-2; Paratype NMNH; 46375/ Casey determ.;elongatus-5/ red square Paralectotype; Phloeodes ; ♀ ovipennis ; Casey 1907; designated I. A. Foley 2006.

Phloeodes elongatus : LECTOTYPE, here designated: ♂, missing left meso tarsus in NMNH. Kern; Co. Cal.,(ifornia)/ Casey; bequest; 1925/ orange rectangle Type NMNH 46374/ hand written elongatus; Csy./ red square Lectotype; Phloeodes ; ♂ elongatus ; Casey 1907; designated I. A. Foley 2006.

Phloeodes scaber : LECTOTYPE, here designated: ♂, intact in NMNH. Handwritten S. Diego/ Casey; bequest; 1925/ Casey determ.; scaber – 4/ orange rectangle Probably; original; TYPE; of scaber/ red square Lectotype; Phloeodes ; ♀ scaber ; Casey 1907; designated I. A. Foley 2006. Of the 7 specimens identified as Phloeodes scaber by Casey, only 2 are from San Diego, and the one designated the Lectotype is closer to the length of the specimen in the original description.

Phloeodes angustus : LECTOTYPE, here designated: ♀, left antenna missing last two antennomeres in NMNH. S. Cal.(ifornia)/ Morrison/ Casey; bequest; 1925/ red rectangle Neo-; Type NMNH; 46378, hand written angustus; Csy./ red square Lectotype; Phloeodes ; ♀ angustus ; Casey 1907; designated I. A. Foley 2006.

Phloeodes remotus : LECTOTYPE, here designated: ♂, right meta-tarsus missing in NMNH. No locality label/ Casey; bequest; 1925/ Casey determ.; remotus – 4/ red square Lectotype; Phloeodes ; ♂ remotus ; Casey 1907; designated I. A. Foley 2006. This species was differentiated by Casey based on the femoral swelling in the male, this specimen is the only male of the 4 Phloeodes remotus identified by Casey. It is also from an unrecorded locality as the original description states.

Discussion of Casey types: The variation in the Casey types is typical of the species throughout its geographic range. The specimens uniquely identified by Casey in key format show the following variation: Phloeodes remotus is densely clothed throughout with a gray-brown vestiture, the velvety spots of the elytra are not black, but a more reddish-brown; in Phloeodes angustus the body is more elongate and narrower in form, the velvety spots indistinct, the dorsal surface is very unevenly and more sparsely tuberculate with large areas along the middle and lateral third devoid of tubercles; in Phloeodes scaber the prothorax is narrower than the elytra, the pronotum is strongly elevated along the median third, and the slopes of the median elevation on the prothorax are slightly concave and devoid of tubercles; in Phloeodes elongatus the prothorax is elongate, the finer tubercles are aggregated, the arcuate ridges are slightly stronger, the elytral velvety spots are large and distinct; Phloeodes ovipennis shows a paler grayish vestiture, the central velvety spot is slightly arcuate and oblique, the basal short, and the dorsal surface of the head has smaller tubercles.

Notes: Gebien (1936) listed a single Casey species, Phloeodes latipennis , as a synonym of Phloeodes pustulosus (LeConte) . This synonymy was not recognized by García-París et al. (2006) who synonymized P. scaber , P. angustus , and P. remotus with P. pustulosus , as well as reporting the Gebien synonymy as new. The remaining Casey species, P. ovipennis and P. elongatus , were also synonymized, but with P. diabolicus . The species described by Casey (1907a, b) are all represented by individuals that are well within the normal character range of Phloeodes diabolicus LeConte. For example the type series of Phloeodes scaber Casey contains seven specimens all of which are a fairly uniform chalky gray color, with the velvety spots on the dorsum of the elytra difficult to see. This is a common situation in specimens where environmental debris from the activity of the insect accumulates on the surface and obstructs some of the external structures. While this series appears slightly different from the other specimens in the Casey collection, it is clearly not a unique and distinct species across the spectrum of conditions that can be found in unprepared specimens of Phloeodes diabolicus . García-París et al. (2006) did not examine any type specimens of Phloeodes ; therefore the opportunity to designate lectotypes is taken here.

García-París et al. (2006) retained P. diabolicus and P. pustulosus as independent taxa, and while they redescribed P. diabolicus , they provided no characters to distinguish between the two species. When LeConte (1859) described P. pustulosus he stated that it was of the “size and form” of Phloeodes diabolicus , but based his discrimination of the two types based on smaller and less regular tubercles, the lack of velvety spots on the elytra, and sculpture near the scutellum. These characters are all clearly variable, and do not justify the division of the continuous population into more than one true species. Clinal variation is seen in the presence ( Fig. 36 ) or absence ( Fig. 35 ) of the whitish vestiture on the elytral humeri, apical declivity, and on the apical ven- trites. Most northern California populations show the whitish vestiture while central and southern populations are uniformly black or dark grey. A similar pattern is seen in the distribution of Zopherus nodulosus nodulosus Solier and Zopherus nodulosus haldemani Horn. Populations north of the Rio Grande River in Texas are predominately white, and those farther south in Mexico have a greater proportion of the dorsal surface black ( Triplehorn 1972).

Figures

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

FIGURES 35–38. Dorsal habiti. 35. Phloeodes diabolicus, California, San Diego Co.; 36. Phloeodes diabolicus, California, Napa Co.; 37. P. plicatus, California, Los Angeles Co.; 38. P. venustus, Costa Rica.

FIGURES 39–41. 39–40. Ventral surface of pronotum. 39. Phloeodes diabolicus; 40. P. plicatus. 41. Lateral view of P. diabolicus hypomeron.

FIGURES 42–46. 42. Lateral view of P. venustus hypomeron. 43. Ventral view of tarsus of P. venustus. 44–46. Male genitalia. 44. P. diabolicus; 45. P. plicatus; 46. P. venustus.

FIGURES 47–49. 47–48. Labial palp insertions. 47. Phloeodes diabolicus; 48. P. plicatus. 49. SEM of tarsal claw of P. plicatus.

FIGURES 50–51. Femora of male Phloeodes diabolicus. 50. Male nodule. 51. Cuticular pores.