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 ) laticollis ( Drury, 1773 ) ( Figs. 46–48 , 118–125 ) Cerambyx laticollis Drury, 1773 : 134 . Prionus laticollis ; Harris, 1841 : 79 ; 1842: 79; LeConte, 1852b : 177 ; Harris, 1852 : 84 ; Emmons, 1854 : 115 ; Fitch, 1859 : 845 (biology); Bland, 1861 (distr.); Holmes, 1869a : 231 (biology); 1869b: 51 (biology); Packard, 1869 : 682 ; 1870: 594 (biology); Riley, 1870 : 87 (biology); Riley, 1873 : 56 (biology); Crotch, 1873 : 83 (checklist); Treat, 1874 : 344 (biology); Saunders, 1875 : 29 (biology); Popenoe, 1877 : 32 (distr.); Crotch, 1880 : 83 (checklist); Riley, 1880 : 238 (host); Packard, 1881 : 118 (biology); Lameere, 1884 : cccxxii (distr.); Weed, 1884 : 13 (biology); Leng, 1884 : 57 ; Blanchard, 1887 : 85 ; Packard, 1890 : 52 , 437 (biology); Bruner, 1891 : 195 (biology); Webster, 1892 : 198 (biology); Hopkins, 1893 : 192 (biology); Bruner, 1894 : 154 (biology); Hamilton, 1895 : 337 (distr.); Beutenmüller, 1896 : 74 (host); Ehrmann, 1897 : 170 (distr.); Wickham, 1897 : 83 ; Bruner, 1899: 162 (biology); Lugger, 1899 : 193 (biology); Smith, 1900 : 285 ; Hopkins, 1902 : 60 (biology); Dury, 1902 : 158 (distr.); Bubna, 1902 : 195 (distr.); Ulke, 1903 : 25 (distr.); Young, 1903 : 158 ; Lockhead, 1903 : 111 (biology); Hopkins, 1904 : 37 (biology); Horsfall, 1904 : 37 (biology); Pettit, 1904 : 41 (biology); Laurent, 1905 : 62 ; Fyles, 1905 : 92 (biology); Felt, 1906 : 486 ; Morris, 1908 : 446 (distr.); Smith, 1910 : 324 ; Blatchley, 1910 : 1011 ; Lameere, 1912a : 236 ; Fisher & Kirk, 1912 : 309 (distr.); Lameere, 1913 : 76 (cat.); Craighead, 1915 : 19 (larva); Johnson, 1915 : 314 (distr.); Lockhead, 1919 : 83 , 85, 322; Nicolay, 1919 : 63 (distr.); Lameere, 1919 : 137 ; Leng, 1920 : 266 (cat.); Britton, 1920 : 266 (distr.); Kempers, 1923 : 102 (morphology); Kirk & Knull, 1926 : 21 (distr.); Leonard, 1928 : 433 (distr.); Hatch, 1930 : 26 (distr.); Beaulne, 1932 : 197 (host); Britton, 1933 : 376 (distr.); Goldman, 1933 : 95 (morphology); Easterling, 1934 : 140 (host); Herrick, 1935 : 220 (biology); Doane et al. , 1936 : 165 ; Saalas, 1936 : 33 (morphology); Britton, 1936 : 258 (distr.); 1938: 144 (distr.); Brimley, 1938 : 210 (distr.); Becker, 1942 : 608 (biology); Löding, 1945 : 113 (distr.); Knull, 1946 : 145 (distr.); Sherman, 1946 : 126 (distr.); Craighead, 1950 : 262 (biology); Jaques, 1951 : 251 ; Beal et al. , 1952 : 71 ; Shenefelt & Benjamin, 1955 : 99 (biology); Nishio, 1956 : 242 ; Linsley, 1957 : 8 (syn.); Alexander, 1958 : 49 (distr.); Gibson & Carrillo, 1959 : 117 (distr.); Dillon & Dillon, 1961 : 577 ; Farrar & Kerr, 1968 : 563 (biology); Benham, 1969 : 1331 (larva, nymph); 1970: 1413 (morphology); Payne et al. , 1970 : 3 (biology); Benham, 1971 : 89 (biology); Swan & Papp, 1972 : 442 ; Baker , 1972 : 200 (biology); Gosling, 1973 : 67 (distr.); Kirk & Balsbaugh, 1975 : 96 (distr.); Benham & Farrar, 1976 : 569 (larva); Turnbow & Franklin, 1980 : 338 (distr.); Campbell et al. , 1989 : 55 ; Chemsak et al. , 1992 : 21 (checklist); Yanega, 1996 : 27 ; Vlasak & Vlasakova, 2002 : 204 (distr.); Robimson, 2005: 85; McCorquodale et al. , 2007 : 121 , 127; Barbour et. al. , 2011: 590, 591; Agnello et al. , 2011 : 17 , 18. Prionus ( Prionus ) laticollis ; Casey, 1912 : 234 , 245; Linsley, 1962 : 39 ; MacRae, 1993 : 227 (distr.); Monné & Giesbert, 1994 : 15 (checklist); Monné, 1995: 52 (cat.); Chemsak, 1996: 103; Schiefer, 1998 : 115 (distr.); Peck & Thomas, 1998 : 116 (distr.); Monné & Hovore, 2005 : 20 (checklist); 2006: 19 (checklist); Özdikmen & Turgut, 2009 : 411 ; Bezark & Monné, 2013 : 28 (checklist). Prionus ( Derobrachus ) laticollis ; Drury & Westwood, 1837 : 78 . Prionus ( Prionellus ) laticollis ; Casey, 1924 : 209 ; Gilmour, 1954 : 45 (distr.). Prionus brevicornis Fabricius, 1801 : 260 ; Harris, 1838 : 89 ; Sturm, 1826 : 187 ; 1843: 239; Haldeman, 1847b : 31 ; LeConte, 1852a : 109 ; Chevrolat 1852 : 650 ; Melsheimer, 1853 : 100 (cat.); White, 1853 : 16 ; Bethune, 1868 : 23 (host); Packard, 1869 : 495 ; Smith, 1873 : 346 ; LeConte, 1878 : 126 ; Packard, 1881 : 127 (host); LeConte & Horn, 1883 : 274 ; Clarkson, 1884 : 95 (biology); Packard, 1890 : 481 (host); Zimsen, 1964 : 163 ( types ). Prionus Brevicornis ; Schönherr, 1817 : 339 (syn.). Prionus ( Prionellus ) brevicornis ; Casey, 1924 : 210 . Prionus ( Prionus ) kempi Casey, 1912 : 233 , 244. Prionus ( Prionellus ) kempi ; Casey, 1924 : 211 . Prionus ( Prionus ) laticollis oblongus Casey, 1912 : 234 ; Lingafelter et al. , 2014 : 87 ( type ). Prionus ( Prionellus ) oblongus ; Casey, 1924 : 210 . Prionus ( Prionus ) parvus Casey, 1912 : 234 ; Lingafelter et al. , 2014 : 296 ( type ). Prionus ( Prionellus ) parvus ; Casey, 1924 : 211 . Prionus ( Prionellus ) frosti Casey, 1924 : 210 ; Lingafelter et al. , 2014 : 66 ( type ). Prionus ( Prionellus ) nigrescans Casey, 1924 : 210 ; Lingafelter et al. , 2014 : 106 ( type ). Prionus nigrescans ; Leonard, 1928 : 433 . Prionus ( Prionellus ) densus Casey, 1924 : 211 ; Lingafelter et al. , 2014 : 52 ( type ). Orthosoma cylindricum ; Holmes, 1868 : 19 (error of identification). Male ( Figs. 118–119 ). Head, mandible, scape, pedicel, antennomere III, pronotum, femora and tibiae from darkbrown to black (often with both colors); elytra from light-brown to black (usually dark-brown or black); antennomeres IV–XI dark-brown (gradually lighter toward last antennomere); palpi from brown to dark-brown; pro- meso, and metasternum, metepisterna dark-brown with some areas darker; ventrites brown, distal edge from yellowish to blackish (commonly with different color, mainly on I and V); tarsi dark-brown, often with small areas darker. Head, excluding mandibles, at central area longer than prothorax, moderately elongate behind eyes (distance from posterior ocular edge to prothorax from slightly shorter to slightly longer than greatest length of upper eye lobe). Longitudinal dorsal furrow distinct from clypeus to prothoracic edge; between middle of eyes (sometimes anterior edge of eyes) and clypeus, placed inside a deep sulcus (broader and deeper toward clypeus). Area on each side of longitudinal sulcus moderately finely punctate (often including inside of sulcus); area close to ocular carinae slightly depressed, coarsely, confluently punctate, with short setae; area around longitudinal furrow, between posterior ocular edge and prothorax, with sub-rhombus, impunctate region; area between and around the latter and prothoracic edge, coarsely (sometimes somewhat finely), moderately sparsely punctate, with short, sparse setae; area behind eyes coarsely, confluently punctate, becoming rugose and pubescent toward lower eye lobe. Antennal tubercles sparsely, moderately coarsely punctate on basal half, becoming impunctate toward apex. Postclypeus narrow, laterally flat, coarsely, confluently punctate, with short, sparse setae; centrally glabrous, impunctate or almost so; anterior edge distinctly concave; area behind central region with deep, narrow, transverse sulcus. Anteclypeus shining, glabrous, impunctate. Labrum triangularly excavated centrally; with long, dense setae. Eyes proportionally large; distance between upper eye lobes from 0.7 to 0.8 times length of scape; distance between lower eye lobes from 1.0 to 1.1 times length of scape. Submentum trapezoid, elevated from base toward mentum; surface from coarsely, abundantly punctate to rugose (sometimes slightly transversely vermiculate); with short, sparse setae centrally, gradually longer and more abundant laterally; anterior edge distinctly carinate. Apex of labial palpi nearly attaining basal one-third of maxillary palpomere IV (sometimes about middle). Latero-basal one-third of mandibles depressed. Antennae with 12 segments; attaining base of distal one-third of elytra. Scape slightly surpassing posterior ocular edge, moderately enlarged toward apex; finely (sometimes barely coarsely) sparsely punctate dorsally; on latero-outer face moderately coarsely punctate; on latero-inner face from smooth to very sparsely and finely punctate. Antennomere III ( Fig. 46 ) dorsally from 1.2 to 1.6 times longer than scape, distinctly enlarged toward apex (distal width equal to about 1.8 times basal width); on dorsal view, imbrication distinct, but not very projected ( Fig. 46 ); on ventral view, apex of imbrication slightly emarginated ( Fig. 47 ); basal two-thirds moderately finely, sparsely punctate; distal one-third very densely, finely punctate (sometimes with small areas with coarse punctures). Antennomere IV about as long as 0.7 times III; dorsally densely, finely punctate throughout (sometimes, sparsely punctate close to the apex); imbrication in dorsal view as in III; distinctly projected backward ventrally, not emarginated at apex. Antennomeres V–VI with sculpture and imbrication as in IV. Antennomeres VII–XI dorsally striolate (striae coarser from VIII); in dorsal view, imbrication more projected than in III; in ventral view as in IV. Antennomere XII about as long as XI; not appendiculate or partially divided. Maximum prothoracic width from slightly shorter to equal to elytral base; anterolateral angle projected forward, rounded or truncate toward first lateral tooth; first lateral tooth small, acute, placed close to anterolateral angle; second lateral tooth large, acute, apex usually slightly projected backwards, placed about middle of margin; margin between second tooth and posterolateral angle from straight to convergent (sometimes with a small tooth at middle); posterolateral angle usually obtuse (sometimes forming a acute angle); basal margin sinuous; distal margin almost straight, emarginated or not centrally. Pronotum usually distinctly convex centrally, explanate laterally; callosities absent or nearly so; disc finely (often slightly coarsely), sparsely punctate (sometimes with central area impunctate or nearly so); coarsely, more abundantly punctate laterally; lateral areas, mainly close to the lateral angles, with long, sparse setae (sometimes short and slightly distinct). Prosternum usually shining, very finely, sparsely punctate (sometimes laterally microsculptured and somewhat opaque); with long, moderately abundant setae (mainly laterally). Prosternal process usually with narrow, longitudinal sulcus shallow in middle; with moderately long, not abundant setae laterally. Elytra coarsely, abundantly punctate (rugose appearance); each elytron with two carinae, usually fused at distal third; sutural spine short. Metasternum depressed centrally toward metacoxae; finely, very densely punctate, except on a subtriangular area along distal one-half of metasternal suture which is microsculptured; with long, dense setae throughout. Metepisterna with sculpture and setae as metasternum laterally. Ventrite I with long, moderately abundant setae along basal one-third (sometimes shorter and sparser), including process, usually shorter and sparser toward lateral margins; ventrites I–IV finely, sparsely punctate, with short sparse setae on base (slightly longer on II); ventrite V finely, densely punctate centrally, with moderately long, abundant setae, sparsely punctate and sparsely and shortly setose laterally. Protarsomeres I–III not spined at apex; mesotarsomere I acute at apex (sometimes with very short spine); mesotarsomeres II–III rounded at apex; metatarsomere I, slender, elongate, not flattened, spined at apex, mainly on inner side; metatarsomere II with short spine at apex on both sides (sometimes only acute); metatarsomere III somewhat acute at apex, slightly longer than II–III together. Female ( Figs. 120–124 ). Head, excluding mandibles, from slightly shorter to slightly longer at middle than prothorax. Dorsal sculpture on face of head and area behind eyes finer, sparser than in male. Distance between upper eye lobes from 0.8 to 0.9 times length of scape; distance between lower eye lobes from 1.1 to 1.2 times length of scape. Submentum as in male. Mandible as in male, but distinctly less sloped on basal one-third, between the carina and inner margin. Antennae nearly reaching apex of basal one-third of elytra; scape distinctly slenderer and longer than in male and slightly surpassing posterior ocular edge; antennomere III ( Fig. 48 ) about 0.9 times as long as scape; antennomeres III–XI without imbrications, distinctly projected distally on outer edge after antennomere V; striae on poriferous system usually present only on antennomeres V–XI, but could be present on IV or apex of III. Prothorax as in male, but typically with posterolateral angle projected; pronotal disc finely, sparsely punctate, mainly on distal one-half. Metasternum and metepisterna from glabrous to very, sparsely short pubescent. Dimensions in mm (male/female) . Total length (including mandibles), 24.5–32.5/35.8–39.0; prothoracic length at center, 3.6–5.0/5.5–5.6; largest prothoracic width, 8.1–10.5/12.6–13.0; humeral width, 8.7–9.8/13.5–13.7; elytral length, 16.5–23.3/23.9–24.5. Geographical distribution. Canada [Ontario ( Bethune, 1868 ), Quebec ( Campbell et al. , 1989 .)], USA [Massachusetts ( Harris, 1838 ); New York ( Emmons, 1854 ), New Jersey ( Bland, 1861 ), Illinois ( Holmes, 1868 ), Missouri ( Holmes, 1869b ), Kansas ( Popenoe, 1877 ), Connecticut ( Smith, 1873 ), Nebraska ( Bruner, 1891 ), Ohio ( Webster, 1892 ), West Virginia ( Hopkins, 1893 ), Pennsylvania ( Hamilton, 1895 ), District of Columbia ( Ulke, 1903 ), Michigan ( Pettit, 1904 ), Indiana ( Blatchley, 1910 ), Virginia ( Craighead, 1915 ), Oklahoma ( Hatch, 1930 ), North Caroline ( Brimley, 1938 ), Alabama ( Löding, 1945 ), South Caroline ( Sherman, 1946 ), Maine ( Gilmour, 1954 ), Wisconsin ( Shenefelt & Benjamin, 1955 ), New Hampshire ( Linsley, 1962 ), Vermont ( Linsley, 1962 ), Rhode Island ( Britton, 1936 ), Delaware ( Linsley, 1962 ), Maryland ( Linsley, 1962 ), Kentucky ( Linsley, 1962 ), Tennessee ( Linsley, 1962 ), Georgia ( Linsley, 1962 ), Mississippi ( Linsley, 1962 ), Florida ( Linsley, 1962 ), Minnesota ( Linsley, 1962 ), South Dakota ( Kirk & Balsbaugh, 1975 ), Arkansas (Chemsak, 1996)]. FIGURES 118–121 . Prionus ( Prionus ) laticollis : 118 , male, dorsal habitus; 119 , male, ventral habitus; 120 , female, dorsal habitus; 121 , female, ventral habitus. Types , type localities. Of Cerambyx laticollis : Drury (1773) did not record the sex of the holotype . Based on the figure from Drury (1770) , the specimen is a female, from USA (New York). Specimens from Drury’s Collection are believed to be lost. However, it is possible that the holotype of C. laticollis has survived, and is deposited in some institution. It could have been sold after Drury’s death, or when he had financial problems and was declared bankrupt, around 1777 ( Ferrer et al. , 2004 ). Of Prionus brevicornis ( Figs. 122–125 ): Fabricius (1801) did not record the sex and number of specimens. According to Tavakilian & Chevillotte (2015) : “ Syntypes (2) / ex collection J. C. Fabricius / Zoologisk Museum, Copenhague / America Boreali. ” However, according to Zimsen (1964) the specimens belong to ZMUK: “ Prionus brevicornis Syst. El. II p. 260. 15 “in America boreali ”.—Kiel 2 specimens .” Webpage of ZMUC (zoology.snm.ku.dk) records: “The majority of Fabrician type specimens are housed in ZMUC, including those deriving from Fabricius’ personal collection, which formally belongs to the Zoological Museum of the University of Kiel ( Germany ), but which is on permanent loan to ZMUC.” Of Prionus ( Prionus ) kempi : Holotype female from USA (New York, Adirondack Mountains), deposited at USNM. Figured at Lingafelter et al. (2016) . Of Prionus ( Prionus ) laticollis oblongus : Described based on males and females. All specimens are from USA (Indiana) and are deposited at USNM. Figured at Lingafelter et al. (2016) . Of Prionus ( Prionus ) parvus : Holotype male “of unknown origin and unindicated locality”, deposited at USNM. Figured at Lingafelter et al. (2016) . Of Prionus ( Prionellus ) frosti : Described based on one male and one female from USA (Massachusetts, Framingham), deposited at USNM. Lingafelter et al. (2014) designated lectotype . Lectotype figured at Lingafelter et al. (2016) . Of Prionus ( Prionellus ) nigrescans : Described on syntypes males from USA (New York, Adirondacks), deposited at USNM. Lingafelter et al. (2014) designated lectotype . Lectotype figured at Lingafelter et al. (2016) . Of Prionus ( Prionellus ) densus : Holotype male from “Locality unrecorded”, deposited at USNM. Figured at Lingafelter et al. (2016) . Material examined. USA , Pennsylvania : Chadds Ford (Delaware County), 1 male , VI.29.1991, J. E. Wappes col. ( MZSP ); 1 female , VII.15 –21.1991, J. E. Wappes col. ( MZSP ). Rhode Island , Lincoln Woods State Park (Providence County), 1 female , VII.19.1997, Cognato col. ( ESSIG ). New Jersey : Camden County, 1 male , VII.15.1928, E. J. F. Marx col. ( ESSIG ); 1 female , VIII.7.1932, E. J. F. Marx col. ( ESSIG ). North Caroline : Great Smoky Mountains National Park, 1 male , VIII.21.1993, L. R. C. F. R. col. ( MZSP ); Cherokee County, 1 male , Nantahala National Forest, Hiwassee Dam, on living white oak tree trunk, Gino Nearns, 4 July 2002 ( ENPC ). Massachusetts : Amherst, 1 male , VII.9.1953, F. Pacheco M. col. ( MZSP ). Tennessee : Monroe County, 1 male , Coker Creek, on living tree trunk, Gino Nearns, 19 July 2003 ( ENPC ); Polk County, 1 male , Farner, on living tree trunk, Gino Nearns, 5 July 2002 ( ENPC ). Remarks. Monné (1995) recorded on the original description of Cerambyx laticollis : “ Drury, 1773 : 83, pl. 37, fig. 2, index.” However Cerambyx laticollis was described in 1770 (volume 1), but it was not until 1773 (volume II) that Drury associated the name with the Linnaean system, giving the index to the figures of the first volume. Although the year is correct, because the name appeared in 1773 (ICZN 1957: Opinion 474), the page, plate, and figure are from volume 1 (published in 1770). According to Santos-Silva et al. (2010) : “It has also been noted that the index is not paginated ( Drury & Westwood 1837 ; Gemminger & Harold 1872 ; Gahan 1895; Lameere 1902; mainly Hayek, 1985: 151), however, this is not the case. The index is paginated as “Mm” which corresponds to page “133”. At the time, it was common to identify each page with a number, and for each four pages to include additional identification of the page as a letter (Aa, Bb, Cc, etc).” The name of Cerambyx laticollis appears in the next page after “Mm”. Thus, the page of the original description is “134” (in the volume II), and not in the page “83” (of the volume I). Schönherr (1817) was the first who recognized that Cerambyx laticollis and Prionus brevicornis were the same species, although he had considered the latter as valid: “19. Brevicornis . * Fabr. S. El. II. p. 260. 15. / Cer. laticollis . * Drury Ins. I. p. 83. T. 37. F. 2.” Drury & Westwood (1837) , without explanation, considered Prionus laticollis in Prionus ( Derobrachus ) Audinet-Serville, 1832 , and also considered Prionus brevicornis equal to P. laticollis , but the latter as having priority: “SYN. Cerambyx Laticollis, Drury, App. vol. 2. / Prionus brevicornis, Fabricius Syst. Eleuth. 2. p. 260. 15. Sch. Syn. Ins. 3. 339. Pal. Bauv. [ sic ] Ins. d’Afr. et d’Amer. Col. Pl. 34. f. 3. ” Harris (1841) , apparently following Drury & Westwood (1837) recorded: “Our largest species is the broad-necked Prionus , Prionus laticollis * of Drury, its first describer.—* Prionus brevicornis of Fabricius.” Later, LeConte (1852b) recognize his mistake in LeConte (1852a) and recorded: “p. 109. Prionus brevicornis . Drury’s Cerambyx laticollis , Vol. 1, pl. 37, fig. 2, has precedence; the name must therefore be Prionus laticollis Harris. Cat. 571.” Between Schönherr (1817) and LeConte (1852b) Prionus brevicornis was listed as valid by some authors who likely considered Prionus laticollis as its synonym: Palisot de Beauvois (1805) , Harris (1838) , Haldeman (1847), Chevrolat (1852) . LeConte (1852a) considered both as the same species, because he made this clear in LeConte (1852b) . However, even after LeConte (1852b) some authors mentioned Prionus brevicornis as having priority (*), distinct of P. laticollis (-), or did not make this clear (+): Melsheimer (1853) (*), White (1853) (*), Bethune (1868) (+), Packard (1869) (-), LeConte (1878) (+), Packard (1881) (+), LeConte & Horn (1883) (+), Clarkson (1884) (+), Packard (1890) (-), and Casey (1924) (-). Lameere (1919) doubted the following species/ subspecies as being synonyms of P. laticollis : “? Kempi Casey … /? oblongus Casey … /? parvus Casey … /? validiceps Casey …” Casey (1924) , without explanation, considered Prionus ( Prionellus ) oblongus as a distinct species of P. ( Prionellus ) laticollis . Linsley (1957) formalized the synonym of Prionus ( Prionus ) kempi , P. ( P. ) laticollis oblongus , P. ( P. ) parvus , synonymized P. ( Prionellus ) frosti , P. ( P. ) nigrescans , and P. ( P. ) densus , and considered P. ( Prionus ) validiceps as synonym of P. pocularis . On Prionus beauvoisi Lameere, 1915 , see remarks on P. imbricornis . According to Linsley (1962) and Chemsak (1996) only the poriferous system of antennomeres VII–XII are striolate. However, all antennomeres are striolate in the males examined by us.