Prionus (Prionus) californicus Motschulsky, 1845

Santos-Silva, Antonio, Nearns, Eugenio H. & Swift, Ian P., 2016, Revision of the American species of the genus Prionus Geoffroy, 1762 (Coleoptera, Cerambycidae, Prioninae, Prionini), Zootaxa 4134 (1), pp. 1-103 : 80-91

Prionus (Prionus) californicus Motschulsky, 1845

( Figs. 111–113 , 158–171, 173–176 , 183 )

Prionus californicus Motschulsky, 1845: 89 ; LeConte, 1852a: 177; Melsheimer, 1853: 100 (cat.); LeConte, 1857: 23 (distr.); 1866: 349 (distr.); Lacordaire, 1868: 61 (note); Gemminger & Harold, 1872: 2757 (cat.); Horn, 1872: 390 (distr.); Snow, 1883: 42 (distr.); Leng, 1884: 57, 58; Rivers, 1886: 64 (biology); Blanchard, 1887: 86; Wickham, 1890: 34 (distr.); Osborn, 1890: 324 (biology); Townsend, 1892: 38; Blaisdell, 1892: 34 (host); Bates, 1892: 144 (distr.); Hamilton, 1894: 30 (distr.); Townsend, 1895: 46 (distr.); Beutenmüller, 1896: 74 (host); Wickham, 1897: 167 (host); Harrington, 1899: 107 (distr.); Wickham, 1899: 123 (distr.); Daggett, 1901: 319 (biology); Fall, 1901: 142 (distr.); Snow, 1906a: 170 (distr.); 1906b: 179 (distr.); Fall & Cockerell, 1907: 191 (distr.); Schaeffer, 1908: 329 (distr.); Lameere, 1912a: 242; 1913: 77 (cat.); Essig, 1915: 251 (biology); Craighead, 1915: 20 (larva); Garnett, 1918: 173 (distr.); Lameere, 1919: 138; Craighead, 1923: 29 (pupa); Essig, 1926: 449 (biology); Tanner, 1927: 33; Hardy & Preece, 1927: 187 (host); Tanner, 1928: 277 (distr.); Crawford & Eyer, 1928: 3 (biology); Knowlton, 1930: 56; Pack, 1930: 219 (distr.); Beaulne, 1932: 219 (host); Barret, 1932:289 (host); Herrick, 1935: 274 (biology); Knowlton & Thatcher, 1936: 278; Doane et al., 1936: 164 (biology); Moore, 1937: 87 (distr.); Quayle, 1938: 319 (biology); Linsley, 1938: 105 (syn.); 1942: 26 (distr.); Hardy, 1942: 10 (biology); Leech, 1947: 141 (biology); Schuh & Mote, 1948: 103 (biology); Balazuc, 1948: 170 (teratology); Knowlton & Wood, 1950: 10 (distr.); Craighead, 1950: 262 (biology); Jaques, 1951: 251; DeLeon, 1952: 80 (host); Keen, 1952: 193 (host); Leech, 1955: 52 (biology); Nishio, 1956: 241; Linsley, 1957: 6 (syn.); Papp, 1959: 85; Linsley et al., 1961: 5 (distr.); Tyson, 1970: 298 (distr.); Swan & Papp, 1972: 441; Tanner & Tanner, 1974: 220 (distr.); Kirk & Balsbaugh, 1975: 96 (distr.); Hovore & Giesbert, 1976: 350 (etology); Furniss & Carolin, 1977: 289 (host); Lewis, 1979: 22 (distr.); Bishop et al., 1984: 20 –24 (biology); Mackay et al., 1987: 363 (distr.); Hovore, 1988: 3 (distr.); Chemsak et al., 1992: 20 (checklist); Noguera & Chemsak, 1996: 396 (distr.); Linsley & Chemsak, 1997: 424 (host); Hanks, 1999: 485, 487 (biology); Weissmann & Kondratieff, 1999: 74 (distr.); Monné, 2002: 23 (host); Barbour et al., 2006: 623; Swiecki & Bernhardt, 2006: 66; Rodstein et al., 2009: 590; Maki et al., 2011: 714; Barbour et al., 2011: 588–592; Rodstein et al., 2011: 114; Hart et al., 2013: 134, 139 (distr.).

Prionus Californicus ; Mannerheim, 1852: 364 (distr.); White, 1853: 16; LeConte, 1876: 519, 520 (distr.).

Prionus (Prionus) californicus ; Casey, 1912: 242; Casey, 1924: 216; Linsley, 1962: 41; Hatch, 1971: 93; Monné & Giesbert, 1994: 14 (checklist); Monné, 1995: 50 (cat.); Chemsak, 1996: 106; Heffern, 1998: 6 (distr.); Monné & Hovore, 2005: 19 (checklist); 2006: 18 (checklist); Monné, 2006: 84 (cat.); Özdikmen & Turgut, 2009: 410; Bezark & Monné, 2013: 27 (checklist); Monné, 2015: 173 (cat.).

Prionus pocularis ; Lord, 1866: 311 (error of identification).

Prionus crassicornis LeConte, 1852a: 108 ; White, 1853: 17; Melsheimer, 1853: 100 (cat.); LeConte, 1857: 23 (syn.); Lacordaire, 1868: 61 (note); Linsley (1938): 105 (syn.).

Prionus (Prionus) crassicornis Casey, 1924: 216 (reval.).

Prionus curvatus LeConte, 1859a: 19 ; Lacordaire, 1868: 61; Gemminger & Harold, 1872: 2757 (cat.); Lameere, 1912a: 242 (syn.); Leng, 1920: 266 (reval.); Leng, 1927: 39; Alexander, 1958: 49 (reval.); Linsley, 1962: 43 (syn.).

Prionus (Prionus) curvatus ; Casey, 1912: 240; 1924: 17 (reval.).

Prionus californicus var. curvatus ; Crotch, 1873: 83; 1880: 83.

Prionus horni Lameere, 1912a: 243 ; Lameere, 1913: 77 (cat.); 1919: 138; Linsley, 1935: 161; 1942: 26; 1957: 7; 1962: 43 (syn.); Damoiseau & Cools, 1987: 33 (type).

Prionus (Prionus) angustulus Casey, 1912: 241 ; Lameere, 1919: 138 (syn. in doubt); Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 17 (type).

Prionus (Prionus) ineptus angustulus ; Casey, 1924: 220 (reval.).

Prionus (Prionus) fissifrons Casey, 1912: 243 ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (cat.; syn.); Casey, 1924: 220 (reval.); Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 63 (type).

Prionus (Prionus) spiculosus fissifrons ; Casey, 1924: 218.

Prionus (Prionus) terminalis Casey, 1912: 244 , 245; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (cat.; syn.); Casey, 1924: 220 (reval.); Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 331 (type).

Prionus (Prionus) californicus ineptus Casey, 1912: 242 ; Lingafelter et al., 2014: 35 (type).

Prionus (Prionus) ineptus ; Lameere, 1919: 138 (syn. in doubt); Casey, 1924: 219 (reval.).

Prionus ineptis [sic]; Linsley, 1938: 105 (syn.).

Prionus ineptus ; Leng, 1920: 266 (cat.; syn.).

Prionus (Prionus) texanus Casey, 1912: 243 ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (cat.; syn.); Casey, 1924: 214 (reval.); Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 332 (type).

Prionus (Prionus) ineptus ambiguus Casey, 1924: 219 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 80 (type).

Prionus (Prionus) ambiguus ; Leng, 1927: 40 (cat.).

Prionus (Prionus) ineptus uintanus Casey, 1924: 220 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 80 (type).

Prionus (Prionus) uintanus ; Leng, 1927: 40 (cat.).

Prionus (Prionus) nanus Casey, 1924: 222 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 103 (type).

Prionus (Prionus) nanus ; Leng, 1927: 40 (cat.).

Prionus (Prionus) spaldingi Casey, 1924: 221 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 322 (type).

Prionus (Prionus) spaldingi ; Leng, 1927: 40 (cat.).

Prionus (Prionus) stultus Casey, 1924: 221 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 326 (type).

Prionus (Prionus) stultus ; Leng, 1927: 40 (cat.).

Prionus (Prionus) stultus parvicollis Casey, 1924: 221 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 327 (type).

Prionus (Prionus) parvicollis ; Leng, 1927: 40 (cat.).

Prionus (Prionus) humeralis Casey, 1924: 216 ; Leng, 1927: 40; Lingafelter et al., 2014: 77 (type).

Prionus humeralis ; Linsley, 1938: 105 (syn.).

Prionus (Prionus) consors Casey, 1912: 240 ; Casey, 1924: 214 (reval.); Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 44 (type).

Prionus consors ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (transf. of syn.).

Prionus (Prionus) consors acomanus Casey, 1912: 241 ; Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 44 (type).

Prionus (Prionus) acomanus ; Casey, 1924: 218 (reval. as distinct species).

Prionus acomanus ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (transf. of syn.).

Prionus (Prionus) consors proximans Casey, 1912: 241 ; Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 45 (type).

Prionus (Prionus) acomanus proximans ; Casey, 1924: 218 (reval.).

Prionus proximans ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (transf. of syn.).

Prionus (Prionus) alutaceus Casey, 1912: 237 ; Casey, 1924: 216 (reval.); Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 14 (type).

Prionus alutaceus ; Lameere, 1919: 138 (syn. in doubt with P. heroicus ).

Prionus (Prionus) suspectus Casey, 1924: 215 ; Leng, 1927: 40; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 330 (type).

Prionus (Prionus) californicus ovipennis Casey, 1924: 217 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 35 (type).

Prionus (Prionus) ovipennis ; Leng, 1927: 40.

Prionus (Prionus) californicus punctulatus Casey, 1912: 243 ; Linsley, 1957: 7 (transf. of syn.); Lingafelter et al., 2014: 35 (type).

Prionus punctulatus ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (transf. of syn.).

Prionus (Prionus) scutellaris Casey, 1924: 219 ; Leng, 1927: 40; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 316 (type).

Prionus (Prionus) spiculosus Casey, 1912: 240 ; Casey, 1924: 217 (reval.); Linsley, 1957: 6 (syn.); Lingafelter et al., 2014: 322 (type).

Prionus spiculosus ; Lameere, 1919: 138 (syn. in doubt); Leng, 1920: 266 (transf. of syn.).

Prionus (Prionus) spiculosus coloradensis Casey, 1924: 218 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 322 (type).

Prionus (Prionus) coloradensis ; Leng, 1927: 40.

Prionus (Prionus) serriger Casey, 1924: 215 ; Leng, 1927: 40; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 318 (type).

Prionus (Prionus) californicus compar Casey, 1924: 217 ; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 35 (lectotype).

Prionus (Prionus) compar ; Leng, 1927: 40.

Prionus (Prionus) orbiceps Casey, 1924: 216 ; Leng, 1927: 40; Linsley, 1957: 7 (syn.); Lingafelter et al., 2014: 35 (type).

Prionus (Prionus) solidus Casey, 1912: 238 ; Casey, 1924: 215; Linsley, 1957: 6 (syn.); Lameere, 1919: 138; Lingafelter et al., 2014: 321 (type).

Prionus (Prionus) tristis Casey, 1912: 236 , 244 (Part: only lectotype female and paralectotype male); 1924: 212; Lameere, 1919: 138; Linsley, 1957: 8 (syn.); Lingafelter et al., 2014: 336 (type). Syn. nov.

Prionus (Prionus) heroicus ; Linsley, 1962: 45 (males); Chemsak, 1996: 109 (males).

Prionus (Prionus) validiceps Casey, 1912: 235 , 245; 1924: 214; Linsley, 1957: 8 (syn.); Lingafelter et al., 2014: 339 (type). Syn. nov.

Prionus (Prionus) tumidus Casey, 1912: 235 , 245; 1924: 213; Lameere, 1919: 138; Linsley, 1957: 8 (syn.); Lingafelter et al., 2014: 337 (type). Syn. nov.

Prionus (Prionus) fontinalis Casey, 1924: 221 ; Lameere, 1919: 138; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 65 (type). Syn. nov.

Note. For redescription of male we are using a specimen of the most common form of the species: eyes large, elytra parallel-sided, metatibiae narrow and elongate (see below). As the current concept of the species is highly variable, we are listing the most common variations after the redescription.

Male ( Figs. 163–169 , 173, 174 , 183 ). Integument reddish-brown, except for: mandibles, most of head, scape, pedicel, part of antennomere III, dorsal surface of femora, inferior margins of femora, base and margins of tibiae, and sternite margins of meso- and metathorax dark-brown to black; base of elytra somewhat darker than remaining surface; metathoracic sternites and abdominal ventrites orange; antennomeres lighter from III to XII. Head, excluding mandibles, about as long as prothorax at central area, slightly elongate behind eyes (distance from posterior ocular edge to the prothorax equal to 0.45 times length of upper eye lobe). Longitudinal dorsal furrow distinct from clypeus to near anterior margin of prothorax, less conspicuous between eyes and prothorax. Frons short, smooth, shining, glabrous. Area on each side of longitudinal furrow, between antennal tubercles and middle of eyes, deeply sulcate, coarsely, sparsely punctate between antennal tubercles, confluently punctate between eyes. Area close to ocular carinae coarsely, confluently punctate, with short, sparse setae. Vertex coarsely, abundantly punctate close to eyes and toward sides, gradually finely, sparsely punctate toward central area close to prothorax; with very short, moderately sparse setae at center, gradually longer toward and behind eyes. Area behind upper eye lobes moderately coarsely, abundantly punctate (punctures coarser toward eye); with short, moderately sparse setae. Area behind lower eye lobes finely rugose, with short, moderately sparse setae near prothorax, with brush of moderately short setae close to eye. Antennal tubercles at base narrowly separated; coarsely, densely punctate at base, finely, sparsely punctate toward apex which is smooth. Postclypeus narrow, moderately finely, abundantly punctate laterally, gradually sparser toward center; with short, moderately abundant setae laterally, gradually sparser toward center. Anteclypeus shining, glabrous, impunctate. Labrum with very long, abundant setae. Eyes moderately large; upper eye lobes wide (about wide as 0.65 times length of scape); distance between upper eye lobes 0.5 times length of scape; distance between lower eye lobes 0.60 times length of scape; distance between posterior ocular edge to prothorax 0.6 times width of upper eye lobe. Submentum trapezoid, slightly depressed, rugose, with short, sparse setae (narrower or wider depending on distance between lower eye lobes); anterior edge distinctly carinate. Apex of labial palpi nearly attaining apex of maxillary palpomere III. Antennae with 12 segments; nearly attaining base of distal one-third of elytra. Scape not attaining posterior ocular edge; moderately enlarged toward apex; moderately finely, abundantly punctate dorsally. Antennomere III 1.15 times as long as scape dorsally, distinctly enlarged toward apex (distal width 2.1 times basal width); on dorsal view, imbrication distinct, projected; on ventral view imbrication with two distinct lobes centrally, separated by deep emargination, with outermost largest, at level of carina not emarginated, finely, sparsely punctate dorsally on basal three-fourths, microsculptured on distal one-fourth. Remaining antennomeres dorsally microsculptured. Antennomere IV 0.65 times as long as III. Imbrication of antennomeres IV–XI as in III, but typically with emargination of outermost lobe distinct.

Maximum prothoracic width about 0.85 times as wide as elytral base; anterolateral angle spined (spine long), anterior margin slightly rounded; lateral spine very long, projected backwards; posterolateral angle acute, projected. Pronotum somewhat flat centrally; disc finely, moderately abundantly punctate, distinctly coarser, denser laterally; with sparse, long setae laterally. Prosternum finely, abundantly punctate, somewhat rugose laterally; with long, moderately dense setae. Elytra finely, abundantly punctate; each elytron with three carinae; sutural apex with short spine.

Ventrites finely, moderately abundantly punctate; with moderately short, sparse setae (notably denser on basal center of I). Pro- and mesotarsomeres I–II acute at apex; apex of lobes of metatarsomere III with short, distinct spine.

Female ( Figs. 111–113 , 158–160, 162 , 170-171, 175–176 ). Integument variable as in male. Head, excluding mandibles, slightly shorter than prothorax at middle. Dorsal sculpture of face on head and area behind eyes finer, sparser than in male. Distance between upper eye lobes 0.65 times length of scape; distance between lower eye lobes 0.65 times length of scape. Submentum as in male. Antennae with 12 segments, nearly reaching middle of elytra; scape distinctly slenderer than in male, not attaining posterior ocular edge; antennomere III 1.35 times as long as scape; imbrication of antennomeres III–XI distinct dorsally at outer side, especially after V. Prothorax as in male; pronotal sculpture as in male. Metasternum and metepisterna with, abundant, long setae throughout.

Variations. Male: body primarily dark-brown to black except; head from reddish-brown to black; elytra mostly orangish-brown; meso- and metathoracic sternites mostly dark-brown; abdominal sternites mostly darkbrown with apex from orange to brown; head at central area from 1.0 to 1.2 times as long as prothorax; head from slightly elongate behind eyes to clearly elongate (distance from posterior ocular edge to prothorax from 0.45 to 1.2 times length of upper eye lobe); frons moderately large; longitudinal dorsal furrow distinct from clypeus to anterior margin of prothorax; Area on each side of longitudinal furrow, between antennal tubercles and middle of eyes, from deeply to slightly sulcate, from moderately coarsely to distinctly coarsely punctate, with punctures between antennal tubercles sparse to abundant, confluently punctate or not between eyes; sculpture on vertex variable, but usually finer and sparser toward center; center of vertex glabrous; antennal tubercles close at base; upper eye lobes from 0.50 to 0.65 times as wide as length of scape; distance between upper eye lobes from 0.50 to 0.75 times length of scape; distance between lower eye lobes from 0.60 to 1.30 times length of scape; distance between posterior ocular edge to prothorax from 0.6 to 1.5 times width of upper eye lobe; antennae attaining base at distal one-fourth of elytra; scape ending from near middle of upper eye lobe to posterior ocular edge; scape slightly enlarged toward apex; punctures on dorsal surface of scape sparse; antennomere III dorsally from 1.15 to 1.40 times as long as scape; distal width of scape from 1.80 to 2.35 times basal width; lobes of imbrication of antennomere III highly variable in size; lobes of imbrication of antennomere III separated by emargination with variable appearance (from almost “V-like” to distinctly wider and shallower; outermost lobe of imbrication of antennomere III at level of carina from not to distinctly emarginated; punctures on dorsal surface of antennomere III present throughout; dorsal side of antennomere IV not totally microsculptured; antennomere IV from 0.60 to 0.75 times length of III; maximum prothoracic width from 0.85 to 0.95 times elytral base; spine of anterolateral angle of prothorax long, with anterior margin from straight to slightly rounded; posterolateral angle of prothorax from acute to obtuse, projected or not; center of pronotal disc with punctures from moderately sparse to abundant, but always fine; sutural elytral apex projected; apex of lobes of metatarsomere III with short, distinct spine on both lobes, or only one lobe, or absent in both lobes. Female: head, excluding mandibles, as long as prothorax at middle; dorsal sculpture on face of head and area behind eyes as in male; distance between upper eye lobes from 0.65 to 0.75 times length of scape; distance between lower eye lobes from 0.65 to 0.85 times length of scape; scape not distinctly slenderer than in male; antennomere III from 1.35 to 1.40 times length of scape; metasternum glabrous or nearly so centrally.

Dimensions in mm (male/female). Total length (including mandibles), 24.2–49.2/36.0–49.0; prothoracic length at center, 3.3–6.0/5.2–7.3; widest prothoracic width, 8.9–15.0/12.1–15.0; humeral width, 9.6–17.5/13.1– 18.3; elytral length, 18.5–35.7/27.2–37.6.

Geographical distribution. Canada [British Columbia ( Hardy & Preece 1927)], USA [Alaska ( Motschulsky 1845), Washington (Jyne 1880), Oregon (LeConte, 1851), Idaho ( Wickham 1897), Montana (Chemsak 1996), California ( Motschulsky 1845), Nevada ( Horn 1872), Wyoming ( Linsley 1962), Utah ( Casey 1924), Colorado ( Hamilton 1894), South Dakota (Chemsak 1996), Nebraska (Chemsak 1996), North Dakota (Heffern 1998), Arizona ( Townsend 1895), New Mexico (LeConte 1859), Texas ( Casey 1912), Arkansas (Heffern 1998), Oklahoma ( Alexander 1958), Georgia (new state record)], Mexico [Sonora ( Bates 1892), Barra California ( Linsley 1942)].

Type, type locality. Of Prionus californicus : Description based on an unspecified number of specimens, male and female, from California ( USA) and Sitka (at that time, Russia, currently Alaska, USA). The species has been recorded as being described based on a single specimen [e.g. Linsley (1962), Monné (1995), and Chemsak (1996)]. However, it is clear that the species was described based on more than one specimen. According to Motschulsky (1845) [translation]: “This species is from Sitka and northern California and must not be confused with the species of North America , quoted in the catalog of Comte Dejean [ Prionus brevicornis Fabricius, 1801 ].” Currently, there are two specimens identified as Prionus californicus in ZMUM, from ex-Collection Motschulsky: one male, without type label; and a female, with type label. Both are from California. According to Mikhail Danilevsky (pers. comm.), a big number of Motschusky’s types were destroyed or lost. As the male specimen deposited at ZMUM lacks a type label, it is not possible to be sure if it is part of the original description. To maintain the stability of the species, we designate as lectotype the female specimen ( Figs. 158–160, 162 ) with the following labels ( Fig. 161 ):

Green (Handwritten): Californ [California]

White (Handwritten): Type

Green (Handwritten): Prionus californicus / Motschs [Motschulsky] / California Red: without letters

Red and yellow (Printed; added by us): LECTOTYPE / Prionus californicus

Of Prionus crassicornis : Description based on at least one male and one female from USA (Oregon), deposited at MCZ. Figured at http://www.mcz.harvard.edu/

Of Prionus curvatus : holotype female from USA (New Mexico, Santa Fé), deposited at MCZ. Figured at http:/ /www.mcz.harvard.edu/

Of Prionus horni ( Fig. 111–113 ): One male and two females from USA (Arizona, Prescott). A couple deposited at NHMW, and a female at IRSN.

Of Prionus (Prionus) angustulus : Holotype male from USA (Arizona, Cañon of the Colorado River), deposited at USNM. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) fissifrons : Holotype male from USA (Arizona), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) terminalis : Holotype male from USA (Arizona, Cañon of the Colorado), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) californicus ineptus : Holotype male from USA (California), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) texanus : Holotype male from USA (Texas, near El Paso), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) ineptus ambiguus: Seven syntypes males from USA (Utah, Eureka), deposited at USNM. Lingafelter et al. (2014) designated lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) ineptus uintanus : Description based on at least one male and one female, from USA (Utah, North Fork, Provo Cañon), deposited at USNM. Lingafelter et al. (2014) designated a male as lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) nanus : Holotype male from USA (Utah, Eureka), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) spaldingi : 13 syntypes (7 males e 6 females) from USA (Utah, Eureka; Colorado, Bounder County). Lingafelter et al. (2014) designated a male as lectotype. The type locality is now Eureka in Utah. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) stultus stultus: Three syntypes (male and female, number of each sex unknown) from the United States (Utah, Eureka), deposited at USNM. Lingafelter et al. (2014) designated lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) stultus parvicollis : Holotype male from USA (Utah, Eureka), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) consors : Syntypes (males and females) from USA (Colorado), deposited at USNM. Lingafelter et al. (2014) recorded: consors Casey, 1912: 240 ( Fig. 46 q, r), Holotype. However, according to Casey (1912): “Length (♂) 34.0–39.5, (♀) 37.0–43.0 mm.; width (♂) 14.5–16.2, (♀) 13.8–17.0 mm.; length and width of prothorax (♂, ♀) 5.5 X 10.0 mm.; width of head (♂) 6.8–7.5, (♀) 6.0– 6.8 mm. Colorado (Boulder Co.). Abundant. Male syntype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) acomanus : Holotype female from USA (New Mexico), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) consors proximans : Holotype female from USA (New Mexico), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) alutaceus : Holotype male from USA (Arizona), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) suspectus : Holotype male from USA (Utah), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) californicus ovipennis : Holotype male from USA (Utah), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) scutellaris : Holotype female from USA (Utah), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) spiculosus : Holotype female from USA (Arizona), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) spiculosus coloradensis : Holotype male from USA (Colorado), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) serriger : Holotype male from USA (Utah), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) compar : Syntypes male from USA (Utah), deposited at USNM. Lingafelter et al. (2014) designated lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Prionus) orbiceps : Holotype male from USA (Utah), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) validiceps ( Figs. 173–174 ): Holotype male from USA (“A single example without indication of locality”), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) tumidus ( Fig. 170–171 ): Holotype female from USA (New Mexico), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) solidus ( Fig. 175 ): Holotype female from USA (Colorado), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Prionus) tristis ( Fig. 176 ). See types of Prionus (Prionus) tristis in P. (P.) heroicus .

Of Prionus (Prionus) fontinalis ( Fig. 183 ): Holotype male from USA (New Mexico), deposited at USNM. Figured at Lingafelter et al. (2016).

Material examined. All types deposited at USNM were examined. USA, Texas: Fort Davis (Jeff Davis County), 1 male, 26.VI.1965, A. & M.E. Blanchard col. ( ESSIG); The Woodlands (Montgomery County), 1 male, 10–18.VI.1977, Wappes col. ( MZSP); 1 female, 15.VI.1978, Wappes col. ( MZSP). Georgia: Dekalb County, 1 male, 2.VII.1971 (no collector indicated) ( MZSP). Arizona: Cochise County, 2 males, Huachuca Mountains, Carr Canyon, Streit, 8/4/1993 ( ENPC); 7 males, Chiricahua Mtns, North Fork, 31˚ 58.265N, 109˚ 18.299W, 1781 m, 23- VII-2009, MV light, N.P. Lord collector ( ENPC); 2 males, same except: Pine Canyon, 31˚ 56.895N, 109˚ 20.495W, 1798 m, 22-VII-2009, MV light, N.P. Lord collector ( ENPC); Cave Creek Canyon (Cochise County), 1 male, 15.VII.1960, J. M. Linsley col. ( MZSP); 1 male, 20–25.VII.2012, Wappes & King col. ( MZSP); Hualapai Mountains (Mohave County), 1 male, 3.VII.1976, Cope col. ( MZSP); 1 male, 2.VIII.1993, Okeefe col. ( ESSIG); Portal, 1 male, 1.VII.1963, Raske col. ( MZSP); 1 male, 9.VII.1963, Raske col. ( MZSP); 5 miles SW Portal female, 25.VII.1960, J. M. Linsely col. ( MZSP); Madera Canyon (Santa Cruz County), 1 female, 30.VI.1988, Stackwell col. ( ESSIG); up Madera Canyon (Madera, 5,600’, 31º42’38”N / 110º52’32”W), 1 male, 18–19.VII.2012, Wappes & King col. ( MZSP); Chiricahua Mountains, 1 male, 9.VII.1959, D.J. & J.N. Knull col. ( MZSP); 1 male, 16.VII.1959, D.J. & J.N. Knull col. ( MZSP); 2 males, 30.VII.1959, D.J. & J.N. Knull col. ( MZSP); M., 1 male, 15.VII.1961, D.J. & J.N. Knull col. ( MZSP). California: San Bernardino Co., 2 females, San Gabriel Mtns., 2,000 m, Wrightwood, at light, Beierl, 8/25/1987 ( ENPC); Berkeley, 1 male, 5.VII.1931, McClay col. ( MZSP); Big Bear Valley (San Bernardino County), 1 female, 12.IX.1967, Stone col. ( ESSIG); Big Creek Reserve (University of California Natural Reserve System; Monterey County), 1 male, 6–8.VII.1992, Powell col. ( ESSIG); Del Puerto Canyon (Stanislaus County; 1,200’; Frank Raines Park), 1 female, 19.V.1972, Rogers col. ( ESSIG); 1 male, 19.V.1972, Kuba col. ( ESSIG); (900–1200’), 1 male, 6.VI.1970, Bentzien col. ( ESSIG); 1 male, 6.VI.1970, Schlinger col. ( ESSIG); Fallsdale (San Bernardino County), 1 female, 21.VII.1956, Johnson col. ( ESSIG); Blodgett Forest Research Station (El Dorado County, 13 miles E Georgewtown), 1 male, 5.VIII.1988, Lindgren col. ( ESSIG); Hat Creek (Shasta County, USFS Work Center), 1 male, VII.1991, Tupy col. ( ESSIG); 1 female, 23.VII.1991, Tupy col. ( ESSIG); Hemet (Riverside County), 1 male, 13.V.1972, Fleming col. ( ESSIG); Kernville (Kern County), 1 male, 1 female, 17.VI.1965, Johnson col. ( ESSIG); Kilkare Woods (Alameda County), 1 male, VII.1921, Hagen col. ( ESSIG); Lone Pine (Owens Valley, Inyo County), 1 male, 8.VI.1937, Johnson col. ( ESSIG); Lytle Creek (San Bernardino County), 1 male, 19.VII.1934, Johnson col. ( ESSIG); 1 male, 13.VIII.1934, Johnson col. ( ESSIG); Miami Creek (Madera County, 4500’), 1 male, 20.VII.1968, O’Brien col. ( ESSIG); Mount Saint Helena, 2 males, V.1930, Linsley col. ( MZSP); Oak Glen (Riverside County), 1 male, 6.VII.1965, Johnson col. ( ESSIG); Oak Knoll Campground (Lake Henshaw, San Diego County), 1 female, VII.1971, Orsak col. ( ESSIG); Orinda (Contra Costa County), 1 male, 16.VII.1985, Hart col. ( ESSIG); Pine Cove (Riverside County, 6100’, San Jacinto Mountains), 2 males, 17.VIII.1986, Brown col. ( ESSIG); Placerville, 1 male, 10.IX.1937, (no collector indicated) ( ESSIG); San Jose, 1 female, 28.VII.1958, Mewaldt col. ( ESSIG); San Mateo, 1 male, 23.VII.1952, DeNoble col. ( ESSIG); Sequoia National Park, 1 male, 15.VII.1972, Whitman col. ( ESSIG); (Whitakers Forest), 1 male, 24.VII.1974, Whitman col. ( ESSIG); Ventura, 1 male, 23.VI.1941, Stone col. ( ESSIG); Waddell (Santa Cruz County), 1 female, 10.VII.1930 ( MZSP); Wildrose, 1 male, VII.1935, Johnson col. ( ESSIG); Yosemite National Park, 1 female, 4.II.1934 (no collector indicated) ( MZSP). New Mexico: Sandoval County, 1 male, Valles Caldera Nat. Pres., Redondo Mdws, 8100 ft. elev., @ UV & MV light, 35˚ 51.70N, 106˚ 36.17W, Nearns & Lingafelter 13- VII-2007 ( ENPC); Las Cruces, 5 males, 1 female, 30.VI.1941, Dawson col. ( ESSIG). Nevada: Leidy Creek ( Esmeralda County , 6200’), 1 male, 7.VII.1965, Bechtel col. ( ESSIG); McIntyre Summit (Nye County, 6940’), 1 male, 23.VII.1964, Bechtel col. ( ESSIG); Peavine Canyon (Nye County, 6300’), 1 male, 9.VIII.1966, Cooney col. ( ESSIG); Peavine Ranch (Nye County), 5 males, 1 female, 29.VII.1964, Cooney col. ( ESSIG); Silver Springs (Lyon County), 1 male, 25.VI.1966, Cooney col. ( ESSIG); 5 miles W Hot Creek Ranch (Nye County), 1 male, 23.VII.1964, Bechtel col. ( ESSIG). Washington: Friday Harbor, 1 male, 7.VIII.1939, Dawson col. ( ESSIG). Utah: Castle Valley (Grand County, 1550 m), 1 male, 12–14.VII.1993, Powell col. ( ESSIG). Colorado: Hesperus, 1 female, 28.VIII.1948, Dawson col. ( ESSIG). MEXICO, Sonora: Rancho Los Alisos, 9.4 km WSW Aconchi, 29º79’83”N / 110º31’97”W, 3 males, 1–2.VII.2013, T. Van Devender col. ( ACMT).

Remarks. Linsley (1942) wrote: “This species has not been previously recorded from Lower California. Three males were taken by Michelbacher and Ross, one of which differs in having thirteen segmented antennae, the last segment of the maxillary palpi widest at the middle, the tarsi more slender, and the elytra rougher. Thus this specimen is even more extreme than the form to which Lameere gave the name Prionus hornii [sic]. Although the antennae are thirteen segmented, the form is not at all like P. lecontei Lameere , from California, which is larger and has very long antennal processes. However, practically all of the forms (species, subspecies, varieties?) which have been named from the californicus complex are based upon males, and until females can be definitely associated with them their status cannot be satisfactorily determined. In the opinion of the present writer they will probably prove to be no more than individual variants of a single plastic species.” We cannot be sure without examination of the specimen, but we believe that the specimen with 13-segmented antennae is P. mexicanus , a species very similar in general appearance to P. californicus .

The first author who synonymized P. crassicornis with P. californicus was LeConte (1857). This synonymy was apparently lost and not mentioned in the more recent catalogues (e.g. Monné, 1995). LeConte (1859a) also confirmed this synonym: “Resembles nearly P. californicus Motsch. ( crassicornis Lec. ), but…”

Later, Casey (1924), without any explanation, revalidated P. crassicornis . After this, Linsley (1938) was the first who considered the former as a synonym of the latter. We agree with the synonymy proposed by Linsley (1938).

Crotch (1873) considered P. curvatus as a variety of P. californicus . This nomenclatural act does not appear in recent catalogues (e.g. Monné, 1995). Prionus curvatus was synonymized with P. californicus by Lameere (1912). Casey (1912) considered the former as distinct from P. californicus . Leng (1920) listed P. curvatus as a valid species, and considered as its synonyms: P. (P.) spiculosus ; P. (P.) consors ; P. (P.) acomanus ; P. (P.) proximans ; P. (P.) angustulus ; P. (P.) horni ; P. (P.) ineptus ; P. (P.) punctulatus ; P. (P.) fissifrons ; P. (P.) texanus ; P. (P.) terminalis . All those synonyms were not observed in the recent catalogues, and many of them are wrongly attributed exclusively to Linsley (1938, 1957), including Linsley himself. Later, Casey (1924), again ignoring Lameere’s works (1912, 1913, 1919), also considered P. curvatus as a distinct species. Alexander (1958) considered P. curvatus as a valid species. Later, Linsley (1962) once again put P. curvatus under synonymy with P. californicus .

Linsley (1935) wrote on P. horni : “I have a number of Prionus from southern Arizona which agree with Lameere’s description of P. ho r n i. These differ from californicus only in their slightly smaller average size and in having the twelfth segment of the antennae appendiculate and longer than the penultimate segment. It seems probable that these are no more than a subspecies of P. californicus .” Linsley (1942) still did not formalize the synonym of P. ho r n i: “…Thus this specimen [of P. californicus from Baja California] is even more extreme than the form to which Lameere gave the name Prionus hornii [sic].” The same position was maintained in Linsley (1957): “ Prionus horni Lameere also appears to me to be based on a variation of infrequent occurrence. It has been taken with the typical form at Prescott, Arizona, and elsewhere.” It was only in 1962 that Linsley listed P. horni as a synonym of P. californicus . We also examined specimens with the twelfth antennomere more or less divided, but as mentioned by Linsley (1935, 1942, 1957) they are just variations of P. californicus . It should be noted that the synonymy between P. h or n i and P. californicus was proposed by Leng (1920).

Casey (1924) ignored the synonymies by Lameere (1919) and Leng (1920), and considered: Prionus (Prionus) angustulus as a subspecies of P. (P.) ineptus ; P. (P.) fissifrons as subspecies of P. (P.) spiculosus ; P. (P.) ineptus as a species different from P. californicus .

Leng (1927) commented on the species described by Lameere and Casey: “This genus [ Prionus ] has been reviewed by Lameere in 1919 and by Casey in 1924 with contradictory results. Lameere considers lecontei Lmr. valid, curvatus Lec. a synonym / horni Lmr. valid / debilis Csy. a synonym / simplex Csy. doubtful / pubicollis Csy. doubtful / debiliceps Csy. doubtful. Casey affirms the validity of the species he described in 1912…”; and “In view of wide difference of opinion [between Lameere and Casey], especially as to 14087 [ Prionus curvatus in Leng (1920) ], no number is assigned to any of the above [the species described by Casey (1924)].”

Apparently, there are at least three groups of specimens within what is currently accepted as Prionus californicus . It is possible that Prionus californicus is a complex of species that we are unable to separate. The groups of specimens can only be recognized in males. Group 1: eyes large, not notably apart from each other dorsally and ventrally, elytra parallel-sided, metatibiae ( Fig. 168 ) narrow and long; Group 2: eyes small, narrow, notably apart from each other, elytra narrowed toward apex, metatibiae ( Fig. 169 ) wider, not clearly elongate; Group 3: eyes as group 1, metatibiae as group 2, elytra variable. These variations are not geographically isolated, nor related to the size of specimen or specimen color. Other highly variable features are: size and shape of prothorax, pubescence of prosternum, and shape of apex of antennomere III. Linsley (1962) separated P. californicus from P. heroicus (males) based on the shape of pronotum (narrow in P. californicus ; very broad in P. heroicus ), shape of antennomeres (antennae with external processes well developed in P. californicus ; antennae with external processes moderate in P. heroicus ), and color (reddish-brown to piceous-brown in P. californicus ; piceous to black in P. heroicus ). However, as seen above, all these features are variable in P. californicus . For example, we examined brownish specimens with external processes of antennae slightly projected as well as very projected; brownish specimens with prothorax very wide as well as distinctly narrower; blackish specimens with prosternum having very sparse setae, or distinctly abundant. Linsley (1962), for P. h e roi c us, also recorded: “posterior tarsi with lobes of third segment angulate at apex but not spinose”. However, we examined specimens distinctly blackish with both lobes almost rounded at apex (not spined), with distinct spine at apex of outer lobe and unarmed at apex of inner lobe, as well as with a spine at apex of both lobes.

In his synonymy of several species described by Casey (1912, 1924) with P. californicus ¸ Linsley (1957) recorded “After studying a long series of specimens of Prionus californicus from various localities in western North America , I have been unable to recognize any of the numerous forms described by Casey as anything but individual variants.” However, some of this same variation also occurs in males redescribed by Linsley as Prionus heroicus . Linsley (1957) also did not comment on the kind of variations he found in P. californicus .

We consider males of P. he ro i c u s sensu Linsley (1962) as P. californicus . We do not know if Linsley (1962) examined females of P. he ro i c u s, but as the description of the metasternum and number of antennomeres agree with the original description, we believe that females of P. heroicus sensu Linsley (1962) correspond to that species.

As seen above, the “external processes” of antennae in P. californicus is variable. Chemsak (1996) used the same features as Linsley (1962) to separate P. californicus from P. heroicus . That author also illustrated the basal antennomeres of both species to show the difference in the “external processes”. However, any of the blackish specimens studied by us have the antennomeres as in his figure of “ P. heroicus ”. Even the holotype male of P. heroicus has the “external processes” the same as his figure. Thus, we consider males of P. he ro i c u s sensu Chemsak (1996) as P. californicus , and the females as P. heroicus .

Females of P. californicus also show considerable variation in the pubescence of the metasternum: long and dense throughout; distinctly sparser centrally than laterally; moderately short and dense laterally, with central area glabrous.

Linsley (1957) synonymized Prionus (Prionus) validiceps Casey, 1912 with Prionus pocularis Dalman, 1817 . Examination of the holotype of the former revealed that the eyes are notably different from that in the latter. Prionus (P.) validiceps is one of the several forms found in P. (P.) californicus . Thus, we propose the transfer of P. (P.) validiceps from synonymy with P. (P.) pocularis to synonymy with P. (P.) californicus .

Linsley (1957) synonymized Prionus (Prionus) tumidus Casey, 1912 with P. (P.) heroicus Semenov, 1907 . Apparently, the synonymy was based on the pubescence of the metasternum (centrally absent). However, we examined specimens of P. (P.) californicus with the same variation. Furthermore, the apex of meso- and metatibiae is spined at the dorsal margin, as occurs in P. (P.) californicus , while in P. (P.) heroicus the apex is truncate.

Linsley (1957) synonymized Prionus (Prionus) fontinalis Casey, 1924 with P. (P.) heroicus . In our opinion, Prionus (Prionus) fontinalis is just a dark male of P. (P.) californicus .

Regarding P. (P.) tristis , see remarks under P. (P.) heroicus .

Figures

FIGURES 107 – 113. 107 – 110, Prionus (Prionus) batesi: 107, syntype male, dorsal habitus; 108, syntype female, dorsal habitus; 109, labels of the syntype male; 110, labels of the syntype female. 111 – 113, Prionus (Prionus) horni, syntype female from IRSN: 111, dorsal habitus; 112, lateral habitus; 113, labels. Photographs by Noël Mal.

FIGURES 158 – 163. Prionus californicus: 158 – 162, lectotype female: 158, dorsal habitus; 159, ventral habitus; 160, detail, ventral habitus; 161, labels; 162, lateral habitus. 163, male from Motschulsky’s collection. Photographs by Aleksey Gusakov.

FIGURES 164 – 169. Prionus (Prionus) californicus, variation in male: 164 – 166, basal antennomeres; 167 – 169, metatibiae, lateral view.

FIGURES 170 – 178. 170 – 171, Prionus (Prionus) tumidus, holotype female: 170, metasternum; 171, apex of mesotibia and mesotarsus. 172, Prionus (Prionus) tetricus, holotype female, ventral habitus. 173 – 174, Prionus (Prionus) validiceps, holotype male: 173, head, dorsal view; 174, head, lateral view. 175, Prionus (Prionus) solidus, holotype female, ventral habitus. 176 – 178, Prionus (Prionus) tristis: 176, lectotype female, ventral habitus; 177, paralectotype female, dorsal habitus; 178, paralectotype female ventral habitus.

FIGURES 179 – 183. 179 – 180, Prionus (Antennalia) fissicornis parviceps, holotype female: 179, head, fronto-lateral view; 180, head and antennae, ventral view. 181, Prionus (Antennalia) thoracicus, holotype female, head and antennae. 182, Prionus (Antennalia) fissicornis transversus, holotype female, ventral habitus. 183, Prionus (Prionus) fontinalis, holotype male, head, dorsal view.

FIGURES 20 – 51. Antennae. 20 – 23, Prionus (Prionus) pocularis: 20, male, scape, pedicel, antennomeres III – IV, dorsal view; 21, male, antennomeres III – IV, lateral view; 22, male, antennomeres III – IV, ventral view; 23, female, antennomeres IX – XI, ventral view. 24 – 28, P. (P.) poultoni: 24, male, scape, pedicel, antennomeres III – IV, dorsal view; 25, male, antennomeres III – IV, latero-ventral view; 26, male, antennomeres III – V, ventral view; 27, female, scape, pedicel, antennomeres III – IV, dorsal view; 28, female, antennomeres VII – IX, ventral view. 29 – 32, P. (P.) aztecus: 29, male, antennomeres III – IV, ventral view; 30, male, antennomeres III – IV, latero-ventral view; 31, female, antennomers VII – IX, ventral view; 32, male, scape, pedicel, antennomeres III – VIII, dorsal view. 33 – 37, P. (P.) mexicanus: 33, male, scape, pedicel, antennomeres III – VIII, dorsal view; 34, male, antennomeres III – IV, latero-ventral view; 35, male, antennomeres III – IV, ventral view; 36, female, scape, pedicel, antennomeres III – IV, dorsal view; 37, female, antennomeres VII – IX, ventral view. 38 – 41, P. (P.) fissicornis: 38, male, scape, pedicel, antennomeres III – V, dorsal view; 39, male, antennomeres III – V, lateral view; 40, male, antennomeres III – V, ventral view; 41, female, antennomeres VIII – X, ventral view. 42 – 45, P. (P.) imbricornis: 42, female, antennomeres VIII – X, ventral view; 43, male, scape, pedicel, antennomeres III – IV, dorsal view; 44, male, antennomeres III – IV, lateral view; 45, male, antennomeres III – IV, ventral view. 46 – 48, P. (P.) laticollis: 46, male, scape, pedicel, antennomere III, dorsal view; 47, male, antennomeres III – VII, ventral view; 48, female, scape, pedicel, antennomere III, dorsal view. 49 – 51, P. (P.) lecontei, male: 49, scape, pedicel, antennomeres III – IV, dorsal view; 50, antennomeres III – IV, latero-ventral view; 51, antennomeres III – IV, ventral view.