Stenaspis verticalis Audinet-Serville, 1834

Eya, Bryan K., 2021, Recharacterization of Stenaspis Audinet-Serville, 1834 with a new species from Mexico (Coleoptera: Cerambycidae: Cerambycinae: Trachyderini), Insecta Mundi 2021 (851), pp. 1-36 : 21-26

Stenaspis verticalis Audinet-Serville, 1834

( Fig. 103–136 )

Stenaspis verticalis Audinet-Serville 1834: 52 ; Dupont 1838: 51; Castelnau 1840: 419; Guérin-Méneville 1844: 218 (syn.); White 1853: 78 (syn.); Strauch 1861: 127 (cat.); Thomson 1864: 208; Lacordaire 1869: 171; Chenu 1870, 311; Gemminger and Harold 1872: 2967 (cat.); Bates 1880: 76 (dist.), 1885: 321 (dist.); Leng 1886: 62; Townsend 1895: 47 (dist.); Fall and Cockerell 1907: 192 (dist.); Casey 1912: 319; Aurivillius 1912: 458 (cat.); Perkins and Swezey 1924: 51 (host.); Smyth 1934: 117 (biol.); Blackwelder 1946: 589 (cat.); Gibson and Carrillo 1959: 120 (dist.); Linsley 1962: 98; Chemsak et al. 1980: 33 (dist.); Terrón 1992: 288 (dist.); Chemsak et al. 1992: 86 (cat.); Chemsak and Noguera 1993: 64 (dist.); Napp 1994: 279; Monné 1994: 35 (cat.); Monné and Giesbert 1994: 152 (cat.); Noguera and Chemsak 1996: 403 (dist.); Noguera et al. 2002: 624 (dist.); Turnbow et al. 2003: 18 (dist.); Hovore 2006: 374 (dist.); Noguera et al. 2007: 312 (dist.); Maes et al. 2010: 618 (dist.); Garcia Morales et al. 2014: 106 (dist.); Noguera and Gutiérrez 2016: 659 (dist.).

Trachyderes superbus Newman 1838: 493 .

Redescription. Male: Length 17–33 mm. Form large, robust; integument reddish-brown to black; head, pronotum, and appendages at least partly red or bright reddish-orange, elytra metallic blue, green or purple, scutellum

103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120

all black or basal half brownish and apically black. Head with vertex and frons rugulose, irregularly punctate, glabrate; mandibles with apices acute; genae irregularly punctate; sparsely covered with appressed pubescence; antennal tubercles prominent, apices acute; eyes moderately large, finely faceted, upper lobes small, well separated; antennae elongate, 11-segmented, exceeding elytral apices by about four segments, scape conical, coarsely separately to irregularly punctate with very short, depressed hair arising from each puncture; apices of antennomere III–VII slightly enlarged and expanded; antennomeres from apex of III laterally carinate with poriferous area on either side of carina, dorsum of antennomeres III–VI glabrate, nitid, finely, sparsely, irregularly punctate, apices densely punctate and densely clothed with short, black depressed hairs, antennomeres from VII densely clothed with short appressed pubescence; antennomere III longer than I, IV shorter than III, V longer than IV, VI subequal to V, VII subequal to VI, VIII subequal to or shorter than VII, IX shorter than VIII, X shorter than IX, XI longest, appendiculate at apical two-fifth. Pronotum 1.5–1.6 times as broad as long, rounded to broadly angulate at sides; anterior angles inflated, broadly rounded; disc with dorsal calli vague, two in anterior half on either side of middle, and three in basal half, one in middle and one each on either side; area between calli flattened, surface coarsely, irregularly punctate, sparsely covered with whitish, transparent hairs; proepisternum inflated, finely, very densely punctate, sparsely pubescent, and demarcated from prosternum and coarsely punctate pronotal disc; prosternum transversely rugose with finely densely punctate transverse subrectangular impressed area on each side of middle; prosternal intercoxal process nitid, irregularly punctate; mesosternum with surface nitid, anterior half and on sides finely, densely, minutely punctate and densely pubescent with appressed, transparent hairs, posterior half sparsely, irregularly punctate in middle; mesosternal intercoxal process sparsely pubescent on sides; metasternum nitid, sparsely punctate, and sparsely pubescent with long, depressed, whitish hairs, sides occasionally densely pubescent; metepisternum densely, minutely punctate, densely pubescent with whitish, depressed hairs. Elytra about 2.2–2.4 times longer than broad, distinctly margined laterally; disc finely to coarsely punctate. Legs with femora slightly clavate, finely, sparsely to coarsely punctate, surface sparsely clothed with short, depressed hairs, outer margin with short, depressed bristles; tibiae slender, surface glabrate, sparsely to coarsely, densely punctate and sparsely covered with short, depressed hairs, inner margin with a row of short, depressed bristles in apical half. Abdomen glabrate, finely, shallowly and rather sparsely punctate in middle, sparsely pubescent with suberect hairs, sides more finely, densely punctate, densely pubescent with appressed, short hairs; fifth sternite truncate to broadly, shallowly emarginate at apex.

Female: 19–37 mm. Form similar to male, head, pronotum and appendages similar in coloration to male; antennae reaching apical fifth of elytra to attaining apices, scape conical, coarsely, separately punctate to finely, separately punctate; apices of antennomeres III–VI slightly enlarged, VII–XI gradually flattened, apices expanded and angulate; antennomere III longer than I, IV shorter than III, V shorter than III, VI subequal to V, antennomeres from VII–X progressively shorter, XI longer than X and subequal to IX, appendiculate at apical quarter. Pronotum laterally tuberculate in middle; anterior angles with obtuse callus on each side; disc finely, sparsely, irregularly punctate to coarsely, contiguously punctate, sparsely covered with transparent, erect hairs; proepisternum coarsely, irregularly punctate and not clearly demarcated from prosternum and pronotal disc as in male; prosternum transversely rugose and coarsely striate-punctate (without finely punctate transverse subrectangular area found in males). Abdomen similar to male, fifth sternite rounded to truncate, shallowly emarginate or vaguely indented at apex.

Materials examined. USA: Texas: Val Verde Co.: near Langtry, 13 Oct. 1994, on Baccharis Linnaeus (Asteraceae) , D.W. Sundberg (1 male, DJHC). MEXICO: Nuevo Leon: 28 km W. Linares Viejo y La Palma, HWY58, 22 Oct. 2004, B.K. EYA (4 females, BKEC). Nayarit: Volcan Ceboruco, 5–15 km W. Jala, 15 Oct. 2001, R.L. Penrose (1 female, LGBC). Jalisco: Est. Biol. Chamela, 1/ 8 July 1988, J.A. Chemsak (1 male, 1 female, BKEC), 23 Oct. 1995, B.K. EYA (1 female, BKEC); Planta La Meza, 13.8 km N.E. San Gabriel, Hwy 432, 18 Oct., B.K. Eya (1 female, BKEC); Ajijic, 1585 m. 11 Sept. 2012, G. Nogueira (1 male, DJHC), 10/ 15 Oct. 1989, D. Curoe (1 male, DJHC); Volcan de Tequila, 1380 m., 25/ 30 Sept. 2012, R. Cunningham, L. Bezark (1 male, 1 female, LGBC). Michoacan: 8.3 km El Puerto (Jct. to Cotija), 19°49′N, 102°37′W, 1800 m., 18 Oct. 1988, A.D. Mudge (1 female, DJHC). Puebla: 22 km N.W. Jct. 190/ 125, 9 km N.W. Oaxaca border, 14 Oct. 2005, B.K. Eya (1 male, BKEC). Guerrero: 23 km W. Iguala, 11/ 16 Sept. 1982, elev. 1040 m., J.A. Chemsak, J.A. Powell (1 female, BKEC). Oaxaca: 1 mi S.E. Tutla, 6500′, 7 Oct. 1977, J. Powell, J. Chemsak, T. Eichlin, and T. Friedlander (1 male, BKEC); 10 km NNE Huajuapan de Leon, 28 Oct. 1990, A.D. Mudge (3 males, 4 females, DJHC, LGBC), 1 Nov. 1990, A. Mudge (3 males,

LGBC), 1 Nov. 1990, A.D. Mudge (1 male, DJHC) ; 12 km NNE Huajuapan de Leon, 1 Nov. 1990, A.D. Mudge (2 males, LGBC) ; 7 km S. Huajuapan de Leon, 1700 m., 28 Oct. 1990, R.L. Westcott (1 female, LGBC) .

Discussion. According to Linsley (1962: 98), this species exhibits considerable geographical variation in color and sculpturing as described by Bates (1885: 321). Stenaspis verticalis males also show considerable allometric variation in the length of antennomeres depending on the size of specimen, where larger individuals (31–33 mm) have antennomere IV longer than III and antennomeres V–X progressively longer, whereas in smaller specimens (17–21 mm) the length of antennomere IV is shorter than III, V longer than IV, VI subequal to or longer than V, and VII–X progressively shorter or subequal to the previous antennomeres. Antennomere XI in males of S. verticalis is the longest and is distinctly appendiculate as though there is a twelfth antennomere but the integument around the appendicular area is definitely fused. In large males (≥ 30 mm) the finely, densely, punctate transverse subrectangular impressed areas on each side of the prosternum are fused in the middle to form one continuous transverse subrectangular impressed area ( Fig. 17 ), whereas in males smaller than 30 mm the transverse subrectangular areas are well separated ( Fig. 20 , 29 ). According to Casey (1912), the holotype of Stenaspis verticalis verticalis Audinet-Serville, 1834 is from Iguala in Guerrero, Mexico although the specific location is not noted by Audinet-Serville (1834) and Dupont (1834) other than just Mexico as the type locality. There are two subspecies named from the USA: Stenaspis verticalis insignis Casey, 1924 , and Stenaspis verticalis arizonicus Casey, 1912 .

Stenaspis verticalis arizonicus Casey, 1912 (Type locality: Arizona, USA). Stenaspis verticalis: LeConte 1858: 40 (cat.). Stenaspis arizonicus Casey 1912: 318 . Stenaspis verticalis arizonicus Linsley 1962: 99 ; Chemsak et al. 1992: 86 (cat.); Monné and Giesbert 1994: 152 (cat.); Monné 1994: 35 (cat.); Lingafelter et al. 2014: 20 (holotype); Heffern et al. 2018: 746 (biol.).

Materials examined. USA: Arizona: Pima Co. : Baboquivari Mountains, Brown Canyon , 10/ 11 July 2001, 31°46′ 23″N, 111°33′31″W, W. K. Will (1 male, 1 female, LGBC) ; Kitt Peak, 6 Aug. 1994, B.K. Eya (1 female, BKEC), Kitt Peak Obs. Rd. on roadside flower, 18 Sept. 1979, C.Y. Kitayama (1 male, LGBC) ; Lower Madera Canyon near old Missile Base , 10 Aug. 1990, R. Duff (1 female, LGBC) ; Kitt Peak Rd. , 1 Sept 1979, L. Bezark, G. Nishida, C. Kitayama, B. Tilden (1 male, LGBC) ; Madera Canyon, 13 Aug. 1988, D. Ahart (1 male, LGBC). Santa Cruz Co.: Pena Blanca Lake , 26 Aug. 1975, L. Bezark, G. Nishida, C. Kitayama, B. Tilden (1 female, LGBC) .

Discussion. According to Linsley (1962: 99), S. v. arizonicus has the elytra finely and densely punctate, usually green or greenish-blue. The pronotum is black and margined posteriorly with red. The ventral surface is reddishbrown or piceous, rarely with any red on metasternum. According to Casey (1912: 319), the very dense sculpture of the elytra is a striking character that differentiates S. v. arizonicus from S. v. verticalis Audinet-Serville. Examination of specimens from Arizona, which is the northwesternmost range of this species, S. v. arizonicus differs from the other subspecies by the darker, duller integument ( Fig. 103–106 , 146 ) and finely, confluently, vermiculate-punctate elytra ( Fig. 137 ). Moving eastward into Texas the punctures on the elytra and scape become coarser, and the integument of the pronotum is more polished, as in the example from Val Verde County, Texas ( Fig. 107–108 , 138 , 147 ).

Stenaspis verticalis insignis Casey, 1924 (Type locality: Texas, Comal Co., USA). Stenaspis insignis: Casey 1924: 262 ; Vogt 1949: 177 (dist.). Stenaspis verticalis insignis: Linsley 1962: 99 ; Hovore et al. 1987: 297 (dist.); Chemsak et al. 1992: 86 (cat.); Monné and Giesbert 1994: 152 (cat.); Monné 1994: 35 (cat.); Lingafelter et al. 2014: 80 (holotype).

Materials examined. USA: Texas: Bexar Co.: Petrenko Rd. 5 Oct. 1994, D.W. Sundberg (1 male, DJHC) ; Camp Bullis, NW. San Antonio , 25 Sept. 1960, E.L. Smith (1 female, LGBC). Medina Co. : 8 mi. W. Median, 15 Sept. 2013, bait, D.W. Sundberg (1 female, DJHC), 4 Oct. 2013, bait, D.W. Sundberg (1 male, DJHC). Comal Co. New Braunfels, 2 Oct. 1984, D. Sundberg (1 female, DJHC) .

Also tentatively assigned to this subspecies are nine additional specimens from Texas as follows: Jim Wells Co.: Premont, 13 Oct. 1984, D.J. Heffern (1 female, DJHC). San Patricio Co.: 4–5 mi. N. Sinton, 12 Oct. 1985, D.J. Heffern (1 female, DJHC); Sinton Area, 9 Nov. 1985, D.J. Heffern (1 male, DJHC). Cameron Co.: Sabal Palm Grove, 25/ 27 Oct. 1986, Heffern and Brattain (1 male, 1 female, DJHC), Hwy 4, Palmito Hill, 21 Oct. 1989, D.J. Heffern (1 female, DJHC); 16 km W. Boca Chica, 24 Oct. 1990, collected from Baccharis Linnaeus (Asteraceae) ,

L.G. Bezark (1 male, 2 females, LGBC).

121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136

Discussion. Examination of specimens of S. v. insignis from the vicinity of San Antonio, Texas (Bexar, Comal, and Medina Counties ), which is the northeasternmost range of this species, shows that this subspecies differs from S. v. verticalis , and S. v. arizonicus by the following characteristics: (1) striking bright reddish coloration of the integument ( Fig. 109–112 ); (2) pronotum that is polished wholly or predominately bright reddish, coarsely, deeply, irregularly punctate, and frequently with two black spots in the anterior half ( Fig. 148 ); (3) prosternum, meso- and metasternum that are mostly or partly reddish; (4) scape and elytra that are coarsely, contiguously punctate ( Fig. 139 , 148 ); and elytra that are usually metallic bluish. The antennae of both sexes of S. v. insignis are shorter than in the other subspecies, and usually about four-fifths the length of the body in females. According to Casey (1924: 263), other than from the overall color of integument, S. v. verticalis differs from S. v. insignis by the sculpture of the elytra where the punctures are well separated. Moving southward from Comal to San Patricio and Cameron Counties in Texas, and further into Nuevo Leon, Mexico, the antennae of both sexes are longer, and in females almost attaining the apex of the elytra or slightly exceeding elytra ( Fig. 115–116 ). Also, there is a darkening of the integument of sternum, proepisternum and pronotal disc (i.e., area which is encompassing the two calli in the anterior half, the middle callus in posterior half, and the anterior margin), and the base of the scape especially in males ( Fig. 113–120 ). A gradual geographical change in metallic coloration of the elytra from blue to greenish is also noticeable.

In general, further south into Mexico (i.e., Nayarit, Jalisco, Michoacán, Guerrero and Puebla) the punctures on the pronotal disc and scape become shallower, finer and sparser ( Fig. 149–153 ), and the basal half of the elytral discs are more separately punctate ( Fig. 140–144 ). There is an exceptional male specimen from Chamela, Jalisco ( Fig. 123–124 , 150 ) with dull reddish integument which resembles S. v. insignis , however, the pronotal disc is coarsely, shallowly and contiguously punctate ( Fig. 150 ) and the elytral disc is finely, separately punctate ( Fig. 141 ). Stenaspis v. insignis has a polished pronotal disc that is coarsely, deeply and irregularly punctate, and the elytra more coarsely, contiguously punctate ( Fig. 139 ). Moving further southeastward from Jalisco into Michoacán, Guerrero and Puebla the elytra of S. v. verticalis become coarser but separately punctate ( Fig. 142–144 ), and further into Oaxaca the elytra become coarsely, deeply and contiguously punctate ( Fig. 145 ), more consistently aeneous green in coloration, and the basal half of the scape is more consistently reddish in all specimens examined ( Fig. 154 ).

Stenaspis validicornis Casey, 1912 (Type locality: Guerrero, Mexico). Stenaspis validicornis Casey 1912: 319 ; Blackwelder 1946: 589 (cat.); Chemsak et al. 1992: 86 (cat.); Monné and Giesbert 1994: 152 (cat.); Monné 1994: 35 (cat.); Noguera and Chemsak 1996: 403 (dist.); Lingafelter et al. 2014: 339, 369 (lect.).

According to Casey (1912: 319), S. validicornis is one of the various forms of the verticalis group, where the males are defined by differences in color of the integument and punctures on the elytral disc and prosternum. The various forms of the verticalis group examined by Casey are based on male characteristics only. Casey (1924: 263), wrote: “ validicornis may be regarded as a subspecies or variety of verticalis Serv. ” According to Mayr (1969: 41), “subspecies are normally allopatric,” and when several subspecies are reported from the same locality, it strongly indicates an incorrect usage of the term subspecies. Since the type localities of S. v. verticalis and S. v. validicornis are both from Guerrero, according to Casey (1912: 319) in close proximity to each other, S. v. validicornis is considered to be consubspecific with S. v. verticalis . This conclusion is based on the polymorphic nature of S. verticalis as a species, which exhibits a notable geographic variation in color and sculpturing and the extensive range in distribution from the southwestern United States to southeastern Mexico. Stenaspis v. validicornis is in the middle of the geographic range of this species and based on the description provided by Casey (1912) S. v. validicornis does not exhibit enough variation in its form and sculpturing to be considered a separate subspecies. A photograph of S. validicornis holotype ( Bezark 2020, id: 26567) does not show any notable difference in the overall appearance compared to S. verticalis (e.g., S. verticalis , syntype, Bezark 2020, id: 18558).

Figures

Figures 103–120. Dorsal and lateral images of Stenaspis verticalis. 103–104) S. v. arizonicus (Casey 1912), male, 29 mm, Pima Co. AZ, USA. 105–106) S. v. arizonicus, female, 27 mm, Pima Co. AZ, USA. 107–108) Male, 23 mm, Val Verde Co. TX, USA. 109–110) S. v. insignis (Casey 1924), male, 30 mm, Medina Co. TX, USA. 111–112) S. v. insignis, female, 28 mm, Comal Co. TX, USA. 113–114) S. v. insignis, male, 29 mm, Cameron Co. TX, USA. 115–116) S. v. insignis, female, 27 mm, Cameron Co. TX, USA. 117–118) Male, 23 mm, Nuevo Leon, MEX. 119–120) Female, 31 mm, Nuevo Leon, MEX.

Figures 121–136. Dorsal and lateral images of Stenaspis verticalis (cont.) 121–122) Female, 26 mm, Nayarit, MEX. 123–124) Male, 31 mm, Jalisco, MEX. 125–126) Female, 37 mm, Jalisco, MEX. 127–128) Female, 23 mm, Michoacán, MEX. 129–130) Female, 26 mm, Guerrero, MEX. 131–132) Male, 32 mm, Puebla, MEX. 133–134) Male, 27 mm, Oaxaca, MEX. 135–136) Female, 27 mm, Oaxaca, MEX.

Figures 15–23. Pronotal and prosternal characteristic of Stenaspis verticalis: 15–17) S. verticalis, male, Jalisco, MEX with sides of pronotum rounded. 15) Disc coarsely punctate with 5 vague calli (red arrows). 16: 16a) Proepisternum inflated and finely punctate (dark reddish area). 16: 16b) Demarcation of proepisternal finely punctate area from coarsely punctate dorsum of disc. 17: 17a) Subrectangular impressed area. 17: 17b) Vague demarcation dividing proepisternum from the subrectangular impressed area. 18, 20) S. verticalis, male, Nuevo Leon, MEX with side of pronotum with distinct lateral tubercles. 19) S. verticalis, male, Puebla, MEX with proepisternum inflated and finely punctate clearly demarcated from dorsum. 21–22) S. verticalis, female, Guerrero, MEX with side of pronotum distinctly tuberculate. 21: 21a) Anterior angles obtusely tuberculate. 22) Pronotal disc and proepisternum coarsely punctate. 23) S. verticalis, female, Jalisco, MEX with prosternum coarsely punctate.

Figures 24–32. Pronotal and prosternal characteristic of Stenaspis castaneipennis. 24–26) S. castaneipennis, male, Oaxaca, MEX with sides of pronotum rounded, disc coarsely punctate with 5 vague calli. 25) Proepisternum inflated and finely punctate (dark reddish area) clearly demarcated from coarsely punctate dorsum (black area). 27–29) S. castaneipennis, male, Oaxaca, MEX with sides of pronotum with distinct lateral tubercles. 28) Proepisternum inflated and finely punctate (reddish area) clearly demarcated from dorsum (black area). 29: 29a) Subrectangular finely punctate area of prosternum. 29: 29b) Demarcation line dividing proepisternum from finely punctate subrectangular area of prosternum. 30–32) S. castaneipennis, female, Oaxaca, MEX with side of pronotum distinctly tuberculate. 30: 30a) Anterior angles obtusely tuberculate. 31–32) Pronotal disc, proepisternum and prosternum coarsely punctate; proepisternun not demarcated from dorsum and prosternum.

Figures 146–154. Changes in sculpturing of scape and pronotum of Stenaspis verticalis from northwestern USA to southeastern MEX. 146) S. v. arizonicus, male, Pima Co. AZ, USA with darker, duller integument, and more finely punctate scape. 147) Male, Val Verde Co., TX, USA with coarser punctures on scape and pronotum. 148) S. v. insignis, male, Medina Co. TX, USA with coarsely, contiguously punctate scapes, and pronotal disc that is polished, predominately bright reddish, and with two black spots in anterior half. 149) Female, Nayarit, MEX with polished, sparsely, finely punctate scape and pronotum. 150) Male, Jalisco, MEX resembling S. v. insignis but with dull reddish integument and pronotal disc that is coarsely, shallowly and contiguously punctate, and scape separately punctate. 151) Female, Michoacán, MEX with sparsely, finely punctate scape and pronotum. 152) Female, Guerrero, MEX with sparsely, finely punctate scape and coarsely punctate pronotum. 153) Male, Puebla, MEX with sparsely, shallowly punctate scape and pronotal disc. 154) Male, Oaxaca, MEX with more densely punctate pronotal disc and basal half of scape that is more consistently reddish.

Figures 137–145. Variations in elytral sculpturing of Stenaspis verticalis from northwestern USA to southeastern MEX. 137) S. v. arizonicus, male, Pima Co. AZ, USA with finely, confluently, vermiculate-punctate elytra. 138) Male, Val Verde, TX, USA with coarser punctures. 139) S. v. insignis, male, Medina Co. TX, USA with coarsely, contiguously punctate metallic bluish elytra. 140) Female, Nayarit, MEX with separately punctate elytra. 141) Male, Jalisco, MEX with finer, separately punctate elytra. 142) Female, Michoacán, MEX with coarser, separately punctate elytra. 143) Female, Guerrero, MEX with even coarser punctures than female from Michoacán. 144) Male, Puebla, MEX with punctures well separated and sparser. 145) Male, Oaxaca, MEX with much coarser, separately punctate aeneous green elytra.