Revision of the American species of the genus Prionus Geoffroy, 1762 (Coleoptera, Cerambycidae, Prioninae, Prionini) Author Santos-Silva, Antonio Author Nearns, Eugenio H. Author Swift, Ian P. text Zootaxa 2016 4134 1 1 103 journal article 10.11646/zootaxa.4134.1.1 a157eae0-8cb8-4b9a-8ce5-dd7ff7913ba3 1175-5326 399754 92AC0E20-F532-4D21-AE1F-4B056327212F Prionus ( Prionus ) pocularis Dalman, 1817 ( Figs. 20–23 , 139–142 ) Prionus pocularis Dalman, 1817 : 148 ; Sturm, 1843 : 239 ; White, 1853 : 17 ; Melsheimer, 1853 : 100 (cat.); Lacordaire, 1868 : 61 (note); Crotch, 1873 : 83 (checklist); 1880: 83 (checklist); Leng, 1884 : 57 , 58; Beutenmüller, 1896 : 74 (host); Smith, 1900 : 285 ; Ulke, 1903 : 25 (distr.); Evans, 1906 : 99 (distr.); Fall & Cockerell, 1907 : 191 (distr.); Leng, 1910 : 77 (distr.); Blatchley, 1910 : 1012 ; Smith, 1910 : 324 ; Lameere, 1912a : 238 ; Dow, 1913 : 78 ; Lameere, 1913 : 77 (cat.); Frost, 1915 : 209 , 210; Craighead, 1915 : 20 (larva); Nycolay, 1919: 63 (distr.); Lameere, 1919 : 137 ; Britton, 1920 : 266 (distr.); Kirk & Knull, 1926 : 21 (distr.); Leonard, 1928 : 433 (distr.); Doane et al. , 1936 : 165 ; Brimley, 1938 : 210 (distr.); Löding, 1945 : 113 (distr.); Knull, 1946 : 146 ; Craighead, 1950 : 262 (biology); Beal et al. , 1952 : 134 ; Linsley, 1957 : 8 ; Dillon & Dillon, 1961 : 580 ; Gosling, 1973 : 67 (biology); Turnbow & Franklin, 1980 : 338 (distr.); Chemsak et al. , 1992 : 21 (checklist); Browne & Peck, 1996 : 2158 (distr.); Yanega, 1996 : 27 ; Linsley & Chemsak, 1997 : 425 (host); Vlasak & Vlasakova, 2002 : 204 (distr.); Monné & Hovore, 2005 : 20 (checklist); 2006: 19 (checklist); McCorquodale et al. , 2007 : 121 . Prionus ( Prionus ) pocularis ; Casey, 1912 : 238 , 244; 1924: 213; Linsley, 1962 : 41 ; MacRae, 1993 : 227 (distr.); Monné & Giesbert, 1994 : 15 (checklist); Monné, 1995: 53 (cat.); Chemsak, 1996: 105; Schiefer, 1998 : 115 (distr.); Peck & Thomas, 1998 : 116 (distr.); Özdikmen & Turgut, 2009 : 411 ; Bezark & Monné, 2013 : 28 (checklist). Prionus pocularius ; Haldeman, 1847b : 31 (error). Prionus laevigatus Harris, 1837 : 83 ; Haldeman, 1847b : 31 ; LeConte, 1852a : 109 ; White, 1853 : 16 ; Melsheimer, 1853 : 100 (cat.); Bland, 1861 : 93 (distr.); Crotch, 1873 : 83 (syn.); Lacordaire, 1868 : 61 (note); Greene, 1918 : 257 . Prionus elongatus Chevrolat, 1838 : 119 (wrong name to P. laevigatus Harris, 1837 ). Prionus obliquicornis LeConte, 1852a : 108 ; White, 1853 : 17 ; Melsheimer, 1853 : 100 (cat.); Crotch, 1873 : 83 (syn.); Lacordaire, 1868 : 61 (note); Lameere, 1913 : 77 (cat.; syn.). Prionus ( Prionus ) obliquicornis ; Casey, 1912 : 233 , 244; 1924: 213. Prionus curticornis LeConte, 1852a : 109 ; Melsheimer, 1853 : 100 (cat.); White, 1853 : 17 (syn.); Lacordaire, 1868 : 61 (note); Lameere, 1913 : 77 (cat.; syn.). Prionus (Prionus) curticornis ; Casey, 1912 : 233 , 244; 1924: 213. Prionus ( Prionus ) pocularis prolixus Casey, 1912 : 239 ; 1924: 213 (syn.); Lingafelter et al. , 2014 : 301 ( type ). Prionus ( Prionus ) bicolor Casey, 1912 : 239 ; 1924: 213; Linsley, 1957 : 8 (syn.); Lingafelter et al. , 2014 : 27 ( type ). Integument from reddish-brown to brown, normally darker dorsally, primarily on head, pronotum and base of elytra; antennae distinctly lighter toward apex; ventrites distinctly lighter; margins of tibiae and femora black or dark-brown. FIGURES 133–138 . Prionus flohri : 133 , Lectotype male, dorsal habitus; 134 , male, paralectotype male, dorsal habitus; 135 , paralectotype female, dorsal habitus; 136 , labels of the lectotype; 137 , labels of the paralectotype male; 138 , label of the paralectotype female. Photographs by Joachim Willers. FIGURES 139–142 . Prionus ( Prionus ) pocularis : 139 , male, dorsal habitus; 140 , male, ventral habitus; 141 , female, dorsal habitus; 142 , female, ventral habitus. Male ( Figs. 139–140 ). Head, excluding mandibles, from 1.00 to 1.15 times as long at central area as prothorax, elongate behind eyes (distance from posterior ocular edge to the prothorax from 0.55 to 0.70 times greatest length of upper eye lobe). smooth, usually punctate laterally. Area between antennal tubercles and middle of eyes coarsely, abundantly punctate (sometimes punctures sparser); with short, sparse setae (sometimes absent centrally or extremely short throughout). Area from middle of upper eye lobes and anterior margin of prothorax coarsely, sparsely punctate (sometimes centrally smooth), but distinctly denser close to eyes; glabrous, except for short, sparse setae close to eyes. Area behind upper eye lobes coarsely, abundantly punctate; with short, moderately sparse setae. Area behind lower eye lobes moderately finely, densely punctate, somewhat rugose; with moderately long, sparse setae closer to prothorax, forming brush close to eyes. Antennal tubercles coarsely, moderately abundantly punctate on basal one-half, smooth toward apex; glabrous or with short, sparse setae on base. Postclypeus coarsely, densely punctate laterally, gradually finer, sparser toward center. Labrum with very long, abundant setae. Eye proportionally large; distance between upper eye lobes from 0.45 to 0.60 times length of scape; distance between lower eye lobes from 0.3 to 0.6 times length of scape. Submentum trapezoid, distinctly narrowed toward gula, somewhat depressed, with anterior margin narrow, distinctly elevated; surface rugose, with moderately long, sparse setae. Apex of labial palpi nearly attaining middle of maxillary palpomere IV. Mandibles from 0.5 to 0.55 times as long as head. Antennae 12-segmented; reaching from base of distal one-third to distal one-fourth of elytra. Scape reaching or nearly reaching posterior ocular edge; finely, sparsely punctate dorsally, slightly denser punctate laterally, mainly on basal one-third. Antennomere III ( Fig. 20 ) about 1.3 times as long as scape dorsally; enlarged toward apex (widest width from 1.7 to 1.9 times basal width); imbrication distinct, but slightly projected ( Fig. 21, 22 ), with apex slightly emarginated; dorsal surface finely, sparsely punctate on basal two-thirds, densely punctate on distal one-third. Dorsal surface of antennomere IV–VII finely, densely punctate; dorsal surface of remaining antennomeres striate. Imbrication of antennomeres IV–XI as in III. Last antennomere simple. Maximum prothoracic width from 0.85 to 0.95 times elytral base; anterolateral angles from acute to distinctly spined, with anterior margin rounded, somewhat projected forward; side with long spine about middle, usually projected backward; posterolateral angle from subrounded to acute, usually slightly projected. Pronotum finely, moderately abundantly punctate centrally, with smooth area close to base; slightly coarsely and densely punctate laterally; with short, sparse setae laterally. Prosternal process slightly longitudinally sulcate about middle. Elytra coarsely, abundantly punctate, usually somewhat rugose toward apex; each elytron with two distinct carinae dorsally, with carina slightly distinct laterally. Metasternum and metepisterna with long, abundant setae. Ventrite I with short, sparse setae on area of central projection; remaining surface glabrous, except occasionally sparse short setae laterally; ventrites II–IV glabrous, except for short, sparse setae laterally; ventrite V with short, sparse setae laterally and apically. Pro- and mesotarsomeres wide; pro- and mesotarsomeres I–II acute at apex (sometimes only slightly); metatarsomeres distinctly slender, mainly I, with apex of I–II spined. Female ( Figs. 141–142 ). Head, excluding mandibles, from 0.85 to 0.95 times length of prothorax at middle. Sculpture on dorsal surface of head from similar to finer and sparser than in male; area behind eyes as in male. Distance between upper eye lobes from 0.45 to 0.80 times length of scape; distance between lower eye lobes from 0.55 to 1.00 times length of scape. Antennae with 12 segments; reaching from near apex of distal one-third to near middle of elytra; scape distinctly slenderer than males, scape from not reaching to surpassing posterior ocular edge; antennomere III from 0.95 to 1.20 times length of scape; antennomeres ventrally ( Fig. 23 ) as in males, but distinctly slender. Prothorax as in male. Metasternum and metepisterna with abundant long setae, but with area around metasternal suture glabrous or nearly so. Dimensions in mm (male/female) . Total length (including mandibles), 30.8–39.8/38.0–42.3; prothoracic length at center, 4.2–5.5/5.3–6.0; widest prothoracic width, 9.4–11.7/12.7–14.0; humeral width, 10.0–14.4/13.7– 16.0; elytral length, 22.4–28.2/28.3–31.2. Geographical distribution. Canada [Ontario ( Evans, 1906 )], USA [ Georgia ( Dalman, 1817 ), Massachusetts ( Frost, 1915 ), Pennsylvania ( Bland, 1861 ), District of Columbia ( Ulke, 1903 ), New Jersey ( Smith, 1900 ), Indiana ( Blatchley, 1910 ), Mississippi ( Lameere, 1912a ), Michigan ( Lameere, 1912a ), Florida ( Casey, 1912 ), New York ( Nicolay, 1919 ), Connecticut ( Britton, 1920 ), North Carolina ( Brimley, 1938 ) , Alabama ( Löding, 1945 ), Minnesota ( Linsley, 1962 ), Illinois ( Linsley, 1962 ), Virginia ( Linsley, 1962 ), Tennessee ( Linsley, 1962 ), Texas ( Linsley, 1962 ), Arkansas ( Linsley, 1962 ), Louisiana, ( Linsley, 1962 ), South Carolina ( Linsley, 1962 ) , Missouri ( MacRae, 1993 ), Wisconsin ( new state record )]. Lord (1986) recorded Prionus pocularis for British Columbia in Canada . The insect specimens, according to him, were identified by “Mr. Walker.” However, the species that is recorded in that area is P. californicus . Thus, the citation is a mistake. White (1853) was the first to record the species in Canada , but without a precise locality. Fall & Cockerell (1907) recorded P. pocularis for New Mexico (Albuquerque and Santa Fé). However, the species is not known to occur in the state. Thus, the specimens were probably incorrectly identified. The citation is kept in the reference list, until the specimens can be examined to confirm or not the identification. Knull (1946) wrote: “This species should occur in Ohio.” We believe that Prionus pocularis probably occurs in Ohio. However, Knull’s (1946) comment is not a formal record, and no other author has recorded the species in Ohio. Type , type locality. Of Prionus pocularis : Holotype male from USA ( Georgia ), deposited at NHRS . Of Prionus laevigatus : Syntypes (at least, two males and one female), from USA , probably deposited at MCZ and MSB. Monné (1995) recorded: “ Type locality, United States , Massachusetts”. Linsley (1962) and Chemsak (1996) also recorded Massachusetts as the states where the specimens were taken. However, Harris (1837) did not say from where in USA the syntypes originated. We do not know what the source of information was for these authors. According to Harris (1837) : “ P. laevigatus, Harris. Catalogue , p. -. / Halsey’s Collection , No. 227. A male. / Cabinet of the Boston Soc. Nat. Hist. No. 1862, male, No. 1360. Female.” According to Harvard University Library (1982): “In 1831 he [Harris] prepared a catalogue of insects for Edward Hitchcock's Report on the Geology, Mineralogy, Botany and Zoology of Massachusetts.” This correspond to “ P. laevigatus, Harris Catalogue , p. -. ” P. laevigatus in Harris (1833 : 579; 1835: 571) and is a nomen nudum . It is not clear if there was or was not a specimen in Harris’ Collection. Harris (1833) wrote: “The insects enumerate in the preceding list, with the exception of about half a dozen, are contained in my cabinet; and most of them were collected in the vicinity of Boston”; and “A few names, without any authorities annexed will also be found on this list; they are applied to species, which, as far as I can ascertain, do not appear to have been published: these it is my hope soon to make known by means of necessary descriptions.” Thus, it is possible that there was(were) specimen(s) in Harry’s Collection. It is also possible that there is(are) specimen(s) in other collections, because Harris (1833) also wrote that he received specimens from some other collectors: “…I am indebted to my friend the Rev. L. W. Leonard, who has still more increased my obligations to him by his disinterested liberality in sending me even the rare and undescribed insects of which he possessed duplicates.” All specimens of “ P. laevigatus ” examined by Harris as base for his paper (1833) are syntypes , because in 1837 he listed this paper (ICZN, 1999: Article 72.4.1, and 72.4.1.1). Unfortunately, we don’t know if Leonard’s Collection survived or, if it did, where it is deposited. Of Prionus obliquicornis : Holotype male from USA ( Georgia ), deposited at MCZ. Figured at http:// www.mcz.harvard.edu/ Of Prionus curticornis : Holotype female from USA ( Georgia ), deposited at MCZ. Of Prionus ( Prionus ) pocularis prolixus : Holotype female from USA (“Locality unrecorded”), deposited at USNM. Figured at Lingafelter et al. (2016) . Of Prionus ( Prionus ) bicolor : Holotype female from USA (Indiana), deposited at USNM. Figured at Lingafelter et al. (2016) . Material examined. USA , Texas : The Woodlands (Montgomery County), 1 male , VI.10 –18.1977, J. E. Wappes col. ( MZSP ); 1 female , VI.15.1978, J. E. Wappes col. ( MZSP ). Georgia : Athens (Dairy Farm), 1 female , 08.VIII.1994 , [no collector indicated] ( UNESP ); Dekalbe County, 1 male , VII.2.1971, [no collector indicated] ( MZSP ). North Carolina : Gastonia , 1 male , VI.8.1939, R. M. McKenzie col. ( MZSP ); Jacksonville, 1 female , VI.1961 , [no collector indicate] ( MZSP ); 1 male , 1 female , VII.7.1962, [no collector indicated] ( MZSP ). Wisconsin : Eau Claire, 1 male , VII.21.1906, H. H. P. Severin col. ( MZSP ). Alabama : Birmingham, 1 male , VII.1970 , [no collector indicated] ( MZSP ). Florida : Lutz, 1 female , V.18.1916, F. W. Friday col. ( ESSIG ). Remarks. White (1853) , without explanation, synonymized P. curticornis with Prionus pocularis . Crotch (1873) synonymized P. laevigatus and P. obliquicornis with Prionus pocularis . Prionus curticornis was not recorded in Crotch (1873) . Leng (1884) followed Crotch (1873) and added two species as synonyms of Prionus pocularis : “ curticornis , Lec. J. A. P. ser. 2, II, p. 109; obliquicornis , Lec. J. A. P. ser. II, p. 108; laevigatus , Harris, Trans. Soc. Htford 1836 , p. 83, t. I, f. 6; Lec. J. A. P. ser. 2, II, p. 109; denticornis , Sturm, Cat. 1836 , p. 187.” Lacordaire (1868) wrongly listed, nearly all those species synonymized with P. pocularis having them as valid species, except P. laevigatus and P. denticornis . Lameere (1912a) listed P. laevigatus , P. obliquicornis , and P. curticornis as synonyms of Prionus pocularis . Casey (1912) , without comments, considered P. c ur t i c o r ni s and P. obliquicornis as valid species. We consider those revalidations as a valid nomenclatural act, and thus, we consider that it was Lameere (1913) who one more time synonymized those species with P. po c u l a r i s . Casey (1924) , in doubt, listed P. curticornis and P. obliquicornis as synonyms. Lameere (1919) , in doubt, considered Prionus ( Prionus ) pocularis prolixus and Prionus ( Prionus ) bicolor to be synonyms of P. pocularis . Casey (1924) formalized the synonymy of Prionus ( Prionus ) pocularis prolixus , but considered Prionus ( Prionus ) bicolor as a valid species. Linsley (1957) formalized the synonymy of P. ( P. ) bicolor as indicated by Lameere (1919) . Although Leng (1884) has recorded “ denticornis , Sturm, Cat. 1836 , p. 187”, there is no Sturm’s catalogue published in 1836. According to Smithsonian Libraries (http://www.sil.si.edu/DigitalCollections/NHRareBooks/ Sturm/sturm-introduction.htm): “In 1796 Sturm published the catalog of his own insect collection, reproduced here. It also is quite small, only 14 cm . (5 1/ 4 in .) tall. As a result of his work and expanding network of contacts with entomologists and other scientists, his collection grew so rapidly that he issued an enlarged second edition only four years later, in 1800, and eventually a third in 1826 and a fourth in 1843.” We surmise, that Leng (1884) was referring to the second edition of Sturm’s Catalogue (1826). Prionus denticornis Sturm, 1826 is a nomen nudum , “synonymized” with Prionus pocularis by Sturm (1843) .