Prionus (Prionus) imbricornis ( Linnaeus, 1767 )

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 : 39-45

publication ID

https://doi.org/ 10.11646/zootaxa.4134.1.1

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lsid:zoobank.org:pub:92AC0E20-F532-4D21-AE1F-4B056327212F

DOI

https://doi.org/10.5281/zenodo.5066949

persistent identifier

https://treatment.plazi.org/id/03AA87AC-FFE5-672D-FF2C-C7AB2F9B8680

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scientific name

Prionus (Prionus) imbricornis ( Linnaeus, 1767 )
status

 

Prionus (Prionus) imbricornis ( Linnaeus, 1767) View in CoL

( Figs. 42–45 View FIGURES 20 – 51 , 95–98 View FIGURES 95 – 98 , 179–180 View FIGURES 179 – 183. 179 – 180 )

Cerambyx imbricornis Linnaeus, 1767: 622 View in CoL ; Villers, 1789: 224 (note).

Cerambyx (Prionus) imbricornis View in CoL ; Gmelin, 1790: 1815.

Prionus coriarius var. imbricornis View in CoL ; Fabricius, 1781: 206.

Prionus imbricornis View in CoL ; Olivier, 1795: 28 (replaced status); Palisot de Beauvois, 1805: 242; Schönherr, 1817: 340; Lepeletier & Audinet-Serville, 1825: 202; Audinet-Serville, 1832: 192; Burmeister, 1836: 68; Sturm, 1843: 239; Haldeman, 1847b: 31; LeConte, 1852a: 108; White, 1853: 17; Melsheimer, 1853: 100 (cat.); LeConte, 1859a: 48; Lacordaire, 1868: 61 (note); Riley, 1870: 89 (biol.); 1872: 140 (biol.); Horn, 1872: 390 (distr.); Crotch, 1873: 83 (checklist); Putnam, 1876: 172, 173 (distr.); Popenoe, 1877: 33 (distr.); Snow, 1878: 67 (distr.); Schwarz, 1878: 456; Webster, 1879: 20 (etology); Crotch, 1880: 83 (checklist); Riley, 1880: 238 (host); LeConte & Horn, 1883: 274; Leng, 1884: 57, 58; Horn, 1885: 89; 1886: 138; Hamilton, 1886: 112 (larva); Blanchard, 1887: 86; Marten, 1890: 60 (biol.); Riley & Howard, 1891: 407 (biol.); Bruner, 1891: 240 (biol.); Forbes, 1894: 106 (larva); Hamilton, 1895: 337 (distr.); Beutenmüller, 1896: 74 (host); Lugger, 1899: 194 (biol.); Ulke, 1903: 25 (distr.); Hebard, 1903: 261 (biol.); Tucker, 1906: 12 (distr.); Kellogg, 1906: 283; Wickham, 1909: 28 (distr.); Blatchley, 1910: 1012; Leng, 1911: 215 (distr.); Lameere, 1912a: 239; Fisher & Kirk, 1912: 309 (distr.); Lameere, 1913: 77 (cat.); Craighead, 1915: 18 (larva); Lameere, 1919: 137; Lutz, 1921: 338; Craighead, 1923: 29 (larva); Kirk & Knull, 1926: 21 (distr.); Leonard, 1928: 433 (distr.); Ware, 1929: 368 (distr.); Beaulne, 1932: 197 (host); Doane et al., 1936: 165 (distr.); Brimley, 1938: 210 (distr.); Löding, 1945: 113 (distr.); Knull, 1946: 146; Craighead, 1950: 261 (host); Jaques, 1951: 251; Beal et al., 1952: 71; Alexander, 1958: 49 (distr.); Anderson, 1960: 401; Dillon & Dillon, 1961: 580; Frost, 1969: 95 (distr.); Payne et al., 1970: 3; Baker, 1972: 200 (biol.); Swan & Papp, 1972: 442; Kirk & Balsbaugh, 1975: 96 (distr.); Payne et al., 1975: 680 (biol.); Mullins, 1975: 43 (biol.); Stein & Tagestad, 1976: 31; Payne et al., 1976: 9 (biol.); Turnbow & Franklin, 1980: 338 (distr.); White, 1985: 281; Yanega, 1996: 27; Linsley & Chemsak, 1997: 424 (host); Hanks, 1999: 487, 493, 494; Robimson, 2005: 85; Buck et al., 2005: 63; Kelley et al., 2006: 252, 253, 254, 255; Lingafelter, 2007: 18 (key), 138; Barbour et. al., 2011: 590, 591, 592; Hart et al., 2013: 134, 139 (distr.).

Prionus View in CoL ? imbricornis View in CoL ; Chevrolat, 1852: 650.

Prionus (Riponus) imbricornis View in CoL ; Casey, 1912: 248 (key).

Prionus (Neopolyarthron) imbricornis View in CoL ; Casey, 1924: 222 (key); Ortenburger & Hatch, 1926: 146 (distr.); Hatch, 1930: 26; Gilmour, 1954: 45 (distr.); Linsley, 1957: 9; Linsley, 1962: 46; Arnett, 1985: 360; Hovore et al., 1987: 294 (distr.); Chemsak et al., 1992: 21 (checklist); Lingafelter & Horner, 1993: 164 (distr.); McRae, 1993: 227 (distr.); Monné & Giesbert, 1994: 16 (checklist); Monné, 1995: 54 (cat.); Chemsak, 1996: 111; Heffern, 1998a: 6 (distr.); Shiefer, 1998: 115 (distr.); Peck & Thomas, 1998: 116 (distr.); Monné & Hovore, 2005: 21 (checklist); 2006: 20 (checklist); Özdikmen & Turgut, 2009: 410; Bezark & Monné, 2013: 27 (checklist).

Prionus (Riponus) imbricornis mimus Casey, 1912: 248 View in CoL ; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 78 (type).

Prionus (Neopolyarthron) imbricornis mimus View in CoL ; Casey, 1924: 222.

Prionus (Riponus) imbricornis brunneus Casey, 1912: 248 View in CoL ; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 78 (type).

Prionus (Neopolyarthron) imbricornis brunneus View in CoL ; Casey, 1924: 222.

Prionus (Riponus) diversus Casey, 1912: 247 View in CoL ; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 54 (type).

Prionus (Neopolyarthron) diversus View in CoL ; Casey, 1924: 222; Hatch, 1930: 26

Prionus robustus View in CoL ; Brimley, 1938: 210 (distr.);

Prionus (Neopolyarthron) imbricornis diversus View in CoL ; Ortenburger & Hatch, 1926: 146 (distr.).

Prionus imbricornis diversus View in CoL ; Alexander, 1958: 49 (distr.).

Prionus (Riponus) diversus cuneatus Casey, 1912: 247 View in CoL ; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 54 (type).

Prionus (Neopolyarthron) cuneatus View in CoL ; Casey, 1924: 222.

Prionus (Neopolyarthron) robustus Casey, 1924: 222 View in CoL ; Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 310 (type).

Prionus debilis Casey, 1891: 21 View in CoL ; Henshaw, 1895: 24 (cat.); Hamilton, 1896: 164 (syn.); Rice, 1981: 459 (distr.); Leng, 1927: 39; Yanega, 1996: 26; Lingafelter, 2007: 12 (key); 138; Lingafelter et al., 2014: 50 (type). Syn. nov.

Prionus (Riponus) debilis View in CoL ; Casey, 1912: 249.

Prionus (Neopolyarthron) debilis View in CoL ; Casey, 1924: 222; Linsley, 1962: 47 (revalidation); Chemsak et al., 1992: 21; McRae, 1993: 227 (distr.); Monné & Giesbert, 1994: 16 (checklist); Monné, 1995: 54 (cat.); Chemsak, 1996: 112; Heffern, 1998b: 174 (distr.); Peck & Thomas, 1998: 116 (distr.); Monné & Hovore, 2005: 21 (checklist); 2006: 20 (checklist); Özdikmen & Turgut, 2009: 410; Bezark & Monné, 2013: 27 (checklist). Syn. nov.

Prionus imbricornis debilis View in CoL ; Leng (1920): 266 (new status); Brimley, 1938: 210 (distr.).

Prionus imbricornis var. debilis View in CoL ; Knull, 1946: 146.

Prionus simplex View in CoL ; Alexander, 1958: 49 (distr.; error of identification).

Prionus beauvoisi Lameere, 1915: 60 View in CoL ; 1919: 138 (syn.). Syn. nov.

Prionus brevicornis View in CoL ; Palisot de Beauvois, 1805: 216 (not Fabricius, 1801).

Prionus (Antennalia) fissicornis parviceps Casey, 1912: 250 View in CoL ; Lameere, 1919: 138 (syn., in doubt); Linsley, 1957: 9 (syn.); Lingafelter et al., 2014: 62 (type). Syn. nov.

Prionus (Antennalia) parviceps View in CoL ; Casey, 1924: 223.

Integument from reddish to brown, distinctly darker dorsally (frequently dark-brown), mainly on head and pronotum (head always totally darker).

Male ( Figs. 95–96 View FIGURES 95 – 98 ). Head, excluding mandibles, from 0.95 to 1.15 times as long as prothorax at central area, elongate behind eyes (distance from posterior ocular edge to the prothorax slightly less than greatest length of upper eye lobe). Longitudinal dorsal furrow distinct from clypeus to level of posterior edge of upper eye lobes (sometimes distinct, or nearly so, up to anterior edge of prothorax). Frons moderately coarsely, sparsely punctate (sometimes punctures denser or partially confluent); area between antennal tubercles with variable sculpture (from as on frons to distinctly coarser and denser); area between upper eye lobes coarsely, confluently punctate (usually more so close to eyes), but sometimes finer, sparser toward longitudinal furrow; dorsal area between eyes and prothorax moderately finely, sparsely punctate, more so near prothorax (sometimes with central region almost smooth); dorsal area with short, sparse setae (denser near upper eye lobes); area behind upper eye lobes moderately finely, confluently punctate (more so close to eyes), but sometimes sparser, with short, moderately sparse setae; area behind lower eye lobes somewhat rugose about middle, moderately abundantly punctate toward inferior side, with short, sparse setae toward prothorax, with long, moderately abundant setae close to eye. Antennal tubercles moderately finely, sparsely punctate on basal one-half, gradually smoother toward apex; glabrous. Postclypeus coarsely, confluently punctate laterally, centrally smooth or almost so; with moderately abundant setae laterally, glabrous centrally. Anteclypeus not or distinctly separated from postclypeus; smooth, glabrous. Labrum with very long, moderately sparse setae on basal one-half, denser toward apex; brush with long setae along anterior margin. Eyes large; distance between upper eye lobes from 0.35 to 0.65 times length of scape; distance between lower eye lobes from 0.4 to 0.7 times length of scape. Submentum trapezoid, distinctly narrow toward gula, somewhat depressed, with anterior margin narrow, distinctly elevated; surface rugose, with short, sparse setae centrally, and long setae laterally. Apex of labial palpi surpassing middle of maxillary palpomere IV. Mandibles about 0.45 times as long as head; latero-basal one-third depressed. Antennae 17- to 20-segmented; nearly attaining distal one-third of elytra. Scape moderately finely, abundantly punctate dorsally on basal one-third, gradually sparser toward apex (sometimes with smooth area about middle); moderately finely and abundantly punctate laterally; glabrous or with short, sparse setae dorsally on basal one-third. Antennomere III ( Fig. 43 View FIGURES 20 – 51 ) slightly longer dorsally than scape, distinctly enlarged toward apex (largest greatest width from 1.65 to 1.80 times basal width); imbrication very distinct, clearly projected ( Fig. 44 View FIGURES 20 – 51 ); apex of imbrication slightly bifurcate ( Fig. 45 View FIGURES 20 – 51 ), but always with innermost portion much larger than outermost (frequently, difference almost indistinct); dorsal surface finely, sparsely punctate centrally on basal two-thirds, densely punctate on distal one-third (sometimes sparsely punctate). Dorsal surface of remaining antennomeres finely, densely punctate (sometimes partially sparsely punctate on basal onethird of basal antennomeres), gradually striate toward distal antennomeres. Last antennomere complex.

Maximum prothoracic width from 0.8 to 0.9 times elytral base; anterolateral angles rounded, projected forward (sometimes only slightly projected); sides with two distinct acute teeth, one close to anterolateral angle (sometimes almost absent) and another about middle; posterolateral angle from rounded to distinctly acute and projected. Pronotum moderately finely and abundantly punctate, usually slightly less dense laterally; glabrous; center of disc from convex to somewhat flat. Prosternum moderately finely, densely punctate, somewhat rugose laterally; with long, abundant setae laterally; usually almost glabrous centrally. Prosternal process not sulcate; almost glabrous centrally. Elytra moderately coarsely, abundantly punctate; each elytron with three carinae, innermost two most distinct; sutural spine short but distinct (sometimes sutural angle not spined, only slightly projected). Metasternum densely microsculptured; with very long, dense setae. Metepisterna with sculpture and setae as that of metasternum.

Ventrite I with moderately long setae on base, especially on anterior process; ventrites I–IV sparsely, finely punctate, more distinct on distal one-third (sometimes, punctures absent or almost so); ventrites II–IV glabrous or almost so; ventrite V finely, sparsely punctate on basal one-half, denser on distal one-half; ventrite V with short, sparse setae on basal one-half, slightly longer, denser on distal one-half. Tarsomeres I–III moderately slender, mainly metatarsomeres; protarsomeres I–II not or slightly acute at apex, not projected; mesotarsomeres I–II acute and projected at apex; metatarsomeres I–II spined at apex of lobes; tarsomeres I–III with spongy setal pads on ventral surface with distinct longitudinal sulcus centrally, wider on metatarsomeres.

Female ( Figs. 97–98 View FIGURES 95 – 98 ). Head, excluding mandibles, from 0.8 to 0.9 times length of prothorax at middle. Sculpture on dorsal face of head and area behind eyes similar to that in male, but more distinct smooth central area on vertex. Distance between upper eye lobes equal to about 0.7 times length of scape; distance between lower eye lobes equal to about 0.9 times length of scape. Submentum as in male. Antennae with 17–22 segments, nearly reaching basal one-third of elytra; scape similar to that in male; antennomere III from 0.80 to 0.85 times length of scape; antennomeres ( Fig. 42 View FIGURES 20 – 51 ) ventrally carinate after VI–VII, with apex slightly emarginate from about middle of antenna. Prothorax similar to that in male. Metasternum densely microsculptured laterally, gradually finer, sparser toward center; glabrous or with very short, sparse setae near metacoxae. Metepisterna microsculptured, glabrous throughout.

Dimensions in mm (male/female). Total length (including mandibles), 23.8–37.5/36.8–43.3; prothoracic length at center, 3.1–5.5/5.3–5.9; widest prothoracic width, 7.1–12.2/12.1–14.0; humeral width, 8.5–14.4/13.4– 16.1; elytral length, 17.9–27.6/25.7–30.4.

Geographical distribution. Canada [Ontario ( Beaulne, 1932)], United State [South Caroline ( Linnaeus, 1767), North Caroline ( Leng, 1911), Illinois ( Webster, 1879), District of Columbia ( Ulke, 1903), Texas ( Tucker, 1906), Georgia ( Hebard, 1903), Indiana (Casey, 1891), New York ( Casey, 1912), Mississippi (Lameere, 1912), Louisiana ( Lameere, 1912a), Kansas ( LeConte, 1859a), Virginia ( Linsley, 1962), West Virginia ( Craighead, 1915), South Dakota ( Gilmour, 1954), Florida ( Schwarz, 1878), Connecticut ( Linsley, 1962), Nebraska ( Bruner, 1891), Alabama ( Löding, 1945), Arkansas ( Hatch, 1930), Colorado (Chemsak, 1996), Iowa (Putman, 1876), Kentucky ( Linsley, 1962), Maryland ( Linsley, 1962), Minnesota ( Linsley, 1962), Missouri ( Riley, 1870), Montana (Chemsak, 1996), New Hampshire (Heffern, 1998), Ohio ( Knull, 1946), Oklahoma ( Ortenburger & Hatch, 1926), Pennsylvania ( Gronovius, 1764), Tennessee ( Linsley, 1962), New Mexico ( LeConte, 1859a), Delaware ( Linsley, 1962), North Dakota (Chemsak, 1996)], Wisconsin (new state record), and Dominican Republic (Palisot de Beauvois, 1805).

Types, type localities. Of Cerambyx imbricornis: According to Linnaeus (1767): “C. thorace marginato bidentato, corpore ferrugineo, elytris mucronatis, antennis perfoliato imbricates brevioribus”; and “Statura C. coriarii. Antennae subtus imbricate 17 laminis ovato-oblongis.”; “Habitat in Carolina .” Linnaeus (1767) recorded two works where the species was described and/or figured: “ Gron. mus. 529 ” and “ Roes. ins. scar. 2. T. 1. fig. 1 ”. The figure in Rösel (1746) shows a male with anterolateral angles of prothorax very prominent. This feature does not agree with the species currently known as P. imbricornis . However, it is not possible to affirm anything based only on that poor drawing. But we can affirm that the specimen used by Rösel (1746), mentioned by Linnaeus and Gronovius (1764), is a syntype of C. imbricornis (ICZN (1999: Articles 3.2, 12.2 and 72.4.1.1). In the same way, the specimen described by Gronovius (1764) is also a syntype of C. imbricornis . Thus, C. imbricornis was described from “ Carolina ” ( Linnaeus, 1767) and Pennsylvania ( Gronovius, 1764). We do not know where the specimens recorded by Rösel and Gronovius are deposited, or if they survived. According to Deny (1948): “If readers of the twelfth edition of the Systema Naturae (1766–1768) mark well the words “Habitat in Carolina . D. Garden,” they will be impressed by the frequency with which that expression appears in Volumes 1 and 2. Linnaeus’ references were to Dr Alexander Garden who made his home in Charleston, South Carolina , until the outbreak of the American Revolution. It is not possible to know how many specimens Linnaeus (1767) used to describe C. imbricornis . We know that at least a syntype is deposited at LSL. Apparently, there are no types of this species deposited at UUZM, because Wallin (2001) did not list Cerambyx imbricornis in this collection. LSL website (2013) records two specimens from Linnaeus’ collection identified as C. imbricornis . The specimen “LINN 3650”, is certainly not Prionus imbricornis : it is a male of P. c o r i a r i u s ( Linnaeus, 1758) from Darenth (Kent, England). The other specimen, “LINN 3649”, we believe is a syntype of C. imbricornis . The specimen agrees well with the original description, including having 19 antennal segments (scape, pedicel, and 17 imbricate segments, as indicated by Linnaeus). Syntype figured at http://linnean-online.org/linnaeus.html

Of Prionus (Riponus) diversus View in CoL : described based on males (at least two specimens) and a single female. All specimens are from USA (Indiana) and are deposited at USNM. Lingafelter et al. (2014) designated lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Riponus) diversus cuneatus View in CoL : described from at least two males from North Carolina (Southern Pines). The specimens are deposited at USNM. Lingafelter et al. (2014) designated lectotype. Lectotype figured at Lingafelter et al. (2016).

Of Prionus (Riponus) imbricornis brunneus View in CoL : Holotype male, from North Caroline (Southern Pines), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Riponus) imbricornis mimus : Holotype male (type locality unknown), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus (Neopolyarthron) robustus : Holotype female from North Carolina (Southern Pines), deposited at USNM. Figured at Lingafelter et al. (2016).

Of Prionus debilis: Described View in CoL based on seven males from Indiana, Missouri, and Kansas, deposited at USNM. Lingafelter et al. (2014) designated lectotype, with Indiana becoming the type locality. Lectotype figured at Lingafelter et al. (2016).

Of Prionus beauvoisi View in CoL : holotype female from Dominican Republic (Santo Domingo), deposited at MNHN.

Of Prionus (Antennalia) fissicornis parviceps ( Figs. 179–180 View FIGURES 179 – 183. 179 – 180 ): holotype female from USA (Texas), deposited at USNM. Figured at Lingafelter et al. (2016).

Material examined. USA, Georgia: Athens, 1 male, 24.VI.1994, [no collector indicated] ( UNESP); Atlanta, 1 male, VII.01.1965, R. E. Fontaine col. ( ESSIG); 1 male, VII.09.1966, R. E. Fontaine col. ( ESSIG); Tattnall County (3 miles E Hwy 147, near Ohoopee), 1 male, VI.19 –20.1998, Wappes & Morris col. ( MZSP). Missouri: Barnhart (Jefferson County), 1 male, VII.31.1937, E. P. Meiners col. ( ESSIG). South Carolina: Wellford , 1 female, VI.16.1937, R. M. McKenzie. Oklahoma: Catoosa, 1 male, VI.08.1939 [no collector indicated] ( ESSIG). Nebraska: Valentine (Cheery County), 1 male, VII.05.1964, J. W. Johnson col. ( ESSIG). North Dakota: Walcott Dunes, 1 male, VII.17.1975, Doug Scott col. ( ESSIG). South Dakota: Lawrence County (McNenny Fish Hatchery, 11 mi. NW Spearfish, 1 female, VII.21.1953, [no collector indicated] ( ESSIG). Louisiana: Chicot State Park, 1 male, VI.23.1971, [no collector indicated] ( MZSP). Wisconsin: Beloit, 1 male, VII.11.1971, [no collector indicated] ( MZSP).

Remarks. Linsley (1962) and Chemsak (1996) indicated that the antennae in Prionus (Neopolyarthron) have from 15 to 20 segments. All specimens examined by us have at least 17 segments, although it is certainly possible that specimens exist with 15 segments.

According to Linnaeus (1767) [translation]: “ Cerambyx with thorax marginate, bidentate, body ferruginous, elytra mucronate, antennae perfoliate, imbricate, short”; and “Length as in Cerambyx coriairus . Antennae underneath imbricate with 17 segments ovate-oblong.” Linnaeus (1767) recorded two works where the species was described and/or figured: “ Gron. mus. 529 ” and “ Roes. ins. scar. 2. T. 1. fig. 1 ”. As seen above, the figure in Rösel (1746) does not agree with the species currently known as P. imbricornis . However, it is not possible to affirm anything based only on that poor drawing.

Fabricius (1781: 206) considered C. imbricornis as a variety of C. coriarius [translation]: “ Cerambyx imbricornis … Linn. Syst. Nat. 2. 622. 5. It is just a variety. Lives in rotten birch in Europe and North America .” Villers (1789: 224) did not make clear his opinion when he wrote on C. coriarius , but apparently he did not agree with Fabricius [translation]: “Obs. 2. Cerambyx imbricornis Linn. mere variety, according to D. Fab.” Gmelin (1790: 1815) listed C. imbricornis with doubtful status [translation]: “Lives in Carolina , stature of coriarius , is this mere variety?” We consider that Olivier (1795: 66: 28, 66: 29) elevated C. imbricornis to species status, although mistakenly attributed the status of this variety to Linnaeus [translation]: “It is of the shape and size of Prione tanner [ P. cori arius ], which Linnaeus said it could be a variety; it differs in the shape and number of antennal segments, which in Prione tanner are composed of only twelve segments, whereas in the latter [ P. imbricornis ] they are twenty in number..” Those changes on the status of C. imbricornis are not mentioned in catalogues (e.g. Monné 1995).

Hamilton (1896) synonymized Prionus debilis under P. imbricornis : “ Prionus debilis Casey, 1891 , An. N. Y. Acad. Sci. vi, 21. This is a synonym of IMBRICORNIS , being a description of a small and well-known race of that species.” Lameere (1912a) also considered Prionus debilis a synonym of P. imbricornis [translation]: “the small newly hatched male specimens with pale elytra were described as distinct species under the name of P. debilis by Casey.” The synonym was simply ignored by Casey (1924). Brimley (1938), apparently following Leng (1920), considered P. debilis as subspecies of P. imbricornis . Knull (1946), without explanation on the status, recorded P. debilis as a variety of P. imbricornis : “Smaller than imbricornis , less convex, light brown in color.” Linsley (1962), also without explanation on the formal revalidation or on the status, considered P. de b i l i s as a distinct species, condition maintained up to now: “This species is very closely related to imbricornis and the two are sympatric over much of the western part of the range of the latter. P. debilis has a smaller average size, however, is apparently always pale rufotestaceous in color, and differs by having the antennae sparsely, coarsely punctate above.”

According to Casey (1891) P. debilis differs from P. imbricornis by “its much smaller size, narrower, more parallel and less convex form, less chitinized and paler elytra, in the more widely separated eyes and in the vestiture of the hind tarsi.” For Linsley (1962), P. debilis differ from P. imbricornis : “Antennae coarsely, sparsely punctate above; color rufotestaceous”, besides smaller size ( P. imbricornis —“Antennae finely, closely punctate above; color brown, rarely rufotestaceous”, and larger size). Chemsak (1996) recorded the same description for males, and added for females: “Elytra with disk finely punctate, scarcely rugose; color reddish-brown”, for P. debilis ; and “Elytra strongly punctate, rugose; color dark brown to piceous, for P. imbricornis .” Lingafelter (2007) separated P. de b i l i s from P. imbricornis in the couplet 16: “Maximum width of pronotum (including spines) distinctly narrower than elytral base. Color reddish-brown”, for P. debilis ; and “Maximum width of pronotum (including spines) about equal to elytral base. Color light to dark reddish-brown”, for P. imbricornis , P. laticollis , and P. pocularis .”

All these features are variable in P. imbricornis . As for differential characters pointed out by Casey (1891), we can affirm that size is highly variable in all species of Prionus , and usually also in nearly all Prioninae . Thus, it is not a reliable feature to separate the species. Regarding body form, we examined specimens with the body shape somewhat variable. Also, comparing the body shape of holotypes males of P. debilis and P. imbricornis brunneus , it is possible to see that they are very similar. The “less chitinized and paler elytra” suggests a newly emerged specimen, as indicated by Lameere (1912a). The distance between upper eye lobes is slightly variable in males of P. imbricornis , as we could see in the specimens examined (independent of size and/or color). Also, comparing this feature in the holotypes of P. diversus and P. imbricornis mimus (both currently in the synonymy of P. imbricornis ) it is evident that the distance between upper eye lobes are different. Also comparing the syntype of C. imbricornis with the lectotype of Prionus debilis , it is possible to see that there is no difference between distance of upper eye lobes, and the elytral color is nearly the same. According to Casey (1891) on the metatarsus: “posterior tarsi very slender, the under surface densely pubescent only in two small spots at the apices of joints one to three.” This description does not agree with the metatarsus of a paralectotype examined. Actually, the metatarsus is identical to that in P. imbricornis .

For the two differences recorded by Linsley (1962), we examined specimens with different size, color, and with antennomeres sparsely or closely punctate above (mainly on III and IV), showing that those features are variable in P. imbricornis . In the specimens examined, when the dorsal area of antennomeres are finely and closely punctate (microsculptured), the area covered by this punctation is variable, being more evident laterally, and always there is a shining, almost impunctate area dorsally; frequently, there are also coarse punctures mixed laterally and/or dorsally. Specimens were examined that agree very well with the lectotype of P. debilis (shape, size, color, etc.) that have very different kind of sculpture on dorsal surface of antennomeres (mainly on III and IV), being laterally microsculptured or only coarsely, sparsely punctate.

The differences between females of P. imbricornis and P. debilis recorded by Chemsak (1996) are inconsistent as the elytral sculpture is highly variable in males and females of P. imbricornis . The holotype male of P. debilis also has the elytra very coarsely and abundantly punctate while the elytral sculpture of typical P. imbricornis is usually not so evident. Thus, if males of P. de b i l i s have elytral punctation coarser than males of P. imbricornis (or at least, equal), there would be no reason to suppose that the elytral punctation in females is any different. In the key to Prionus (Neopolyarthron) males, Chemsak (1996) figured antennomere III completely finely punctate dorsally. We did not see this condition in the specimens examined, but it is possible that this happens in the species. The antenna figured as being from a male of P. debilis is a female antenna. As in males, the sculpture of female antennae is variable.

The differences recorded by Lingafelter (2007) are also variable in P. imbricornis . Comparing the lectotype of P. diversus with the lectotype of P. diversus cuneatus , and also with the type males of P. imbricornis brunneus and P. i. mimus , it is possible to see just how much the width and shape of the prothorax can vary.

Based on the mix of differential characters for the two species, we place P. de b i l i s as a new synonym of P. imbricornis .

Casey (1924), without explanation, considered Prionus cuneatus , described as subspecies of Prionus diversus , as a distinct species. Ortenburger & Hatch (1926) recorded: “In the list the numbers before the species are those used in C. W. Leng’s “Catalogue of the Coleoptera of America North of Mexico ” (1920).” According to Leng (1920): “In cataloguing the names proposed by Casey in Prionus , the introductory paragraph on p. 215 of his Mem. III, stating that some apply to forms connected by intermediates and destined to speedy extinction, coupled with the suggestion on p. 245 that the abnormal characters employed to distinguish some might not prove constant with more material, has been assumed to warrant doubt to their standing, as to cause the omission of numbers”. Leng (1920) also recorded: “89. imbricornis (L.) 67-622 Ind. Ill.-La. Fla. N.Y. / a. debilis Csy. 91-21 Ind. Mo. Kan. / diversus Csy. 12-247 Ind. / cuneatus Csy. 12-247 B.C. / mimus Csy. 12-248? / brunneus Csy. 12- 248 N.C.” In his catalogue, Leng (1920) used a letter to indicate when he considered a species as subspecies. Thus, he considered Prionus debilis as a subspecies of Prionus imbricornis , disagreeing with Hamilton (1896), although, Leng (1920) was apparently following Lameere (1919), who followed the former. As seen above, Lameere (1919) considered the species listed above by Leng (1920) as possible synonyms, except P. debilis , listed as a true synonym by Lameere (1912, 1913, 1919). Using this concept, and the explanation above on Casey’s species (and subspecies), based on the position in which Leng (1920) included those names (“ diversus ”, “ cuneatus ”, “mimus”, and “brunneus”), it is possible to infer that Leng (1920) was considering them as possible subspecies of P. imbricornis . However, the citations by Leng (1920) are very confused, because “ mimus ” and “brunneus” were described as subspecies of P. imbricornis , and “ cuneatus ” as subspecies of P. debilis . Thus, as it is not possible to know if Leng (1920) was suggesting synonyms or subspecies, it is obligatory to consider without change the condition of the species and subspecies between Lameere (1919) and Linsley (1957), except the status of Prionus debilis that was formally listed as subspecies of Prionus imbricornis . Apparently Ortenburger & Hatch (1926) considered that Leng (1920) changed the status of Prionus diversus to subspecies of P. imbricornis . Alexander (1958) followed Ortenburger & Hatch (1926), recording Prionus imbricornis diversus .

Lameere (1919), in doubt, considered Prionus (Riponus) imbricornis mimus , P. (R.) imbricornis brunneus , P. (R.) diversus , and P. (R.) diversus cuneatus , as synonyms of Prionus imbricornis . However, it was Linsley (1957) who formally established the synonym of those species under P. imbricornis : “I have been unable to detect any population significance for any of the variants named by Casey. Three of his names were based on specimens from Southern Pines, North Carolina (brunneus, cuneatus , robustus ). A fourth was from Indiana ( diversus ). The type locality for mimus was not indicated.”

Lameere (1919) also indicated as synonyms of P. imbricornis , without any explanation: P. beauvoisi Lameere, 1915 , and P. brevicornis sensu Palisot de Beauvois, 1805 (not P. brevicornis Fabricius, 1801 ). Linsley (1957, 1962), and Chemsak et al. (1992) did not include the first as a valid species or as a synonym of any species. Monné & Giesbert (1994), Monné (1995), Monné & Hovore (2005, 2006), and Monné & Bezark (2011) recorded P. beauvoisi as synonym of P. laticollis . We do not know why those latter authors of catalogues and checklists transferred P. b e au v o i s i from the synonym of P. imbricornis to the synonym of P. laticollis . According to Lameere (1915) [translation]: “One female of Prionus from Santo Domingo, with antennae with 14 segments, was described and figured by Palisot de Beauvois (Ins. Afr. et Amer., 1805, p. 216, t. 34, f. 3) under the wrong name of P. brevicornis F. It does not belong to any known species, and I propose to designate this insect, that needs to be found, under the name of P. Beauvoisi .” According to Palisot de Beauvois (1805) [translation]: “ PRIONUS BREVICORNIS . FAB? Piceous; thorax tridentate, dark, shiny; antennae short, 14-segmented, latest 10 antennomeres laterally serrate on one side; elytra punctate ( Fig. 3 View FIGURES 1 – 19 ). Santo Domingo ”; “Obs. This insect has much to do with PRIONUS coriarius . It is much thicker. Although the description of Fabricius is very terse and incomplete, I think that this species is the same that he named BREVICORNIS .”

Prionus beauvoisi View in CoL cannot be P. laticollis View in CoL , because the holotype has antennae with 14 segments, while in the latter the females have antennae with 12 segments. According to Linsley (1962) and Chemsak (1996), females of P. imbricornis View in CoL have antennae with from 15 to 18 segments. However, the holotype female of P. ro bu s t u s, synonymized with P. imbricornis View in CoL by Linsley (1957), has antennae with 19 segments (according to Casey). In turn, the holotype of P. beauvoisi View in CoL has antennae with 14 segments. Thus, both holotype specimens do not agree with the description by Linsley (1957) and Chemsak (1996) on the number of antennal segments. Also, the drawing of P. beauvoisi View in CoL , does not agree with P. imbricornis View in CoL as its prothorax seems different.

Curiously, either P. b e au v o i s i is not mentioned in the synonymic list of P. laticollis View in CoL (or P. imbricornis View in CoL ) or, when it is listed, Dominican Republic is not mentioned as the country where the species occurs ( P. laticollis View in CoL ).

It is not possible to affirm anything about P. beauvoisi without examining the holotype female. If P. beauvoisi really is not P. laticollis , then P. brevicornis sensu Palisot de Beauvois (1805) cannot be listed in the reference list of P. laticollis , because the specimen used as model to the drawing is the holotype of Lameere’s species.

The holotype of Prionus (Antennalia) fissicornis parviceps has the typical antennomeres of P. (P.) imbricornis . Thus, we are transferring the subspecies from the synonymy of P. (P.) fissicornis to that of P. (P.) imbricornis .

In summary, we agree with the synonyms of P. imbricornis established by Linsley (1957), and consider as synonyms of this species: Prionus (Riponus) diversus , P. (R.) diversus cuneatus , P. (R.) imbricornis mimus , P. (R.) imbricornis brunneus , P. (Neopolyarthron) robustus , P. debilis , P. b e au v o i s i, Prionus (Antennalia) fissicornis parviceps , and P. brevicornis sensu Palisot de Beauvois (adding three species and one reference).

MZSP

Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Coleoptera

Family

Cerambycidae

Genus

Prionus

Loc

Prionus (Prionus) imbricornis ( Linnaeus, 1767 )

Santos-Silva, Antonio, Nearns, Eugenio H. & Swift, Ian P. 2016
2016
Loc

Prionus imbricornis diversus

Alexander 1958: 49
1958
Loc

Prionus simplex

Alexander 1958: 49
1958
Loc

Prionus imbricornis var. debilis

Knull 1946: 146
1946
Loc

Prionus robustus

Brimley 1938: 210
1938
Loc

Prionus (Neopolyarthron) imbricornis diversus

Ortenburger 1926: 146
1926
Loc

Prionus (Neopolyarthron) imbricornis

Bezark 2013: 27
Ozdikmen 2009: 410
Monne 2005: 21
Heffern 1998: 6
Shiefer 1998: 115
Peck 1998: 116
Monne 1994: 16
Lingafelter 1993: 164
McRae 1993: 227
Chemsak 1992: 21
Arnett 1985: 360
Linsley 1962: 46
Linsley 1957: 9
Gilmour 1954: 45
Hatch 1930: 26
Ortenburger 1926: 146
Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) imbricornis mimus

Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) imbricornis brunneus

Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) diversus

Hatch 1930: 26
Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) cuneatus

Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) robustus

Lingafelter 2014: 310
Linsley 1957: 9
Casey 1924: 222
1924
Loc

Prionus (Neopolyarthron) debilis

Bezark 2013: 27
Ozdikmen 2009: 410
Monne 2005: 21
Heffern 1998: 174
Peck 1998: 116
Monne 1994: 16
McRae 1993: 227
Chemsak 1992: 21
Linsley 1962: 47
Casey 1924: 222
1924
Loc

Prionus (Antennalia) parviceps

Casey 1924: 223
1924
Loc

Prionus imbricornis debilis

Brimley 1938: 210
Leng 1920: 266
1920
Loc

Prionus beauvoisi

Lameere 1915: 60
1915
Loc

Prionus (Riponus) imbricornis

Casey 1912: 248
1912
Loc

Prionus (Riponus) imbricornis mimus

Lingafelter 2014: 78
Linsley 1957: 9
Casey 1912: 248
1912
Loc

Prionus (Riponus) imbricornis brunneus

Lingafelter 2014: 78
Linsley 1957: 9
Casey 1912: 248
1912
Loc

Prionus (Riponus) diversus

Lingafelter 2014: 54
Linsley 1957: 9
Casey 1912: 247
1912
Loc

Prionus (Riponus) diversus cuneatus

Lingafelter 2014: 54
Linsley 1957: 9
Casey 1912: 247
1912
Loc

Prionus (Riponus) debilis

Casey 1912: 249
1912
Loc

Prionus (Antennalia) fissicornis parviceps

Lingafelter 2014: 62
Linsley 1957: 9
Lameere 1919: 138
Casey 1912: 250
1912
Loc

Prionus debilis

Lingafelter 2014: 50
Lingafelter 2007: 12
Yanega 1996: 26
Rice 1981: 459
Leng 1927: 39
Hamilton 1896: 164
Henshaw 1895: 24
1895
Loc

Prionus

Chevrolat 1852: 650
1852
Loc

Prionus brevicornis

Beauvois 1805: 216
1805
Loc

Prionus imbricornis

Hart 2013: 134
Lingafelter 2007: 18
Kelley 2006: 252
Buck 2005: 63
Hanks 1999: 487
Linsley 1997: 424
Yanega 1996: 27
White 1985: 281
Turnbow 1980: 338
Stein 1976: 31
Payne 1976: 9
Kirk 1975: 96
Payne 1975: 680
Mullins 1975: 43
Baker 1972: 200
Swan 1972: 442
Payne 1970: 3
Frost 1969: 95
Dillon 1961: 580
Anderson 1960: 401
Alexander 1958: 49
Beal 1952: 71
Jaques 1951: 251
Craighead 1950: 261
Knull 1946: 146
Loding 1945: 113
Brimley 1938: 210
Doane 1936: 165
Beaulne 1932: 197
Ware 1929: 368
Leonard 1928: 433
Kirk 1926: 21
Craighead 1923: 29
Lutz 1921: 338
Lameere 1919: 137
Craighead 1915: 18
Lameere 1913: 77
Lameere 1912: 239
Fisher 1912: 309
Leng 1911: 215
Blatchley 1910: 1012
Wickham 1909: 28
Tucker 1906: 12
Kellogg 1906: 283
Ulke 1903: 25
Hebard 1903: 261
Lugger 1899: 194
Beutenmuller 1896: 74
Hamilton 1895: 337
Forbes 1894: 106
Riley 1891: 407
Bruner 1891: 240
Marten 1890: 60
Blanchard 1887: 86
Hamilton 1886: 112
Leng 1884: 57
LeConte 1883: 274
Crotch 1880: 83
Riley 1880: 238
Webster 1879: 20
Snow 1878: 67
Schwarz 1878: 456
Popenoe 1877: 33
Putnam 1876: 172
Crotch 1873: 83
Horn 1872: 390
Riley 1870: 89
Lacordaire 1868: 61
LeConte 1859: 48
White 1853: 17
Melsheimer 1853: 100
LeConte 1852: 108
Haldeman 1847: 31
Sturm 1843: 239
Burmeister 1836: 68
Audinet-Serville 1832: 192
Lepeletier 1825: 202
Schonherr 1817: 340
Beauvois 1805: 242
Olivier 1795: 28
1795
Loc

Prionus coriarius var. imbricornis

Fabricius 1781: 206
1781
Loc

Cerambyx imbricornis

Villers 1789: 224
Linnaeus 1767: 622
1767
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