Pachybrachis subfasciatus (J. E. LeConte, 1824 )

Barney, Robert J., 2017, Definition and Revision of theLuridusSpecies-Group of North AmericanPachybrachisChevrolat (Coleoptera: Chrysomelidae: Cryptocephalinae), The Coleopterists Bulletin 71 (1), pp. 1-45 : 5-22

1. Pachybrachis subfasciatus (J. E. LeConte, 1824) ( Figs. 1, 9A–D, Map 1A)

Cryptocephalus subfasciatus J. E. LeConte 1824: 173 .

Pachybrachys biguttatus Suffrian 1852: 167 .

Pachybrachys impurus Suffrian 1852: 186 . Fall 1915: 377 (taxonomy, catalogue); Knaus 1916 –17: 262 (catalogue). New synonymy.

Pachybrachys xanthias Suffrian 1852: 199 . Fall 1915: 378 (taxonomy, catalogue).

Pachybrachys impurus var. umbrosus Fall 1915: 379 (taxonomy, catalogue). Knaus 1916 –17: 262 (catalogue). New synonymy.

Pachybrachys subfasciatus (J. E. LeConte) : Haldeman 1849: 260 (taxonomy, catalogue); LeConte 1880: 209 (taxonomy, catalogue); Blatchley 1910: 1126 (taxonomy, catalogue); Fall 1915: 376 (taxonomy, catalogue); Johnson 1915: 316 (catalogue, plant association); Lovell 1915: 115 (catalogue, plant association); Dozier 1922: 118 (catalogue); Hughes 1944: 132 (catalogue); Wilcox 1954: 390 (catalogue).

Map 1. Distribution of Pachybrachis species based upon specimens examined. A) P. subfasciatus (black, former

P. impurus grey), and P. dilatatus (red), B) P. femoratus (black) and P. shirleyae (red).

Pachybrachis impurus Suffrian : Barney 1984: 137 (catalogue); Clark et al. 2004: 157 (plant associations).

Pachybrachis subfasciatus (J. E. LeConte) : Melsheimer 1853: 126 (catalogue); Mombert 1869: 544 (catalogue); Downie and Arnett 1966: 1319 (taxonomy); Kirk 1969: 89 (catalogue), 1970: 86 (catalogue); Balsbaugh and Hays 1972: 31 (catalogue); Barney 1984: 144 (catalogue); Riley et al. 2003: 163 (catalogue); Clark et al. 2004: 161 (plant associations); Ciegler 2007: 182 (taxonomy); Barney et al. 2011: 10 (catalogue), 2013: 113 (catalogue).

Type Material. LeConte (1824) described the holotype of P. subfasciatus as inhabiting Georgia .

Fall (1915) apparently did not examine the holotype as no specimens from Georgia were listed in his distribution of specimens. MCZ is not in possession of LeConte’ s holotype; however, LeConte’ s collection at MCZ-LEC consists of a series of five specimens (1♂ 4♀♀) each labeled “[pink circular disc] // P. subfasciatus / (Lec.) Hald. [hand-inked, white paper]”. The pink disc is LeConte’ s color code locality label for “Middle States”, believed to be Connecticut, Delaware, Maryland, New York, New Jersey, Pennsylvania, and Rhode Island. Because none of these specimens are from the holotype locality, the only male available from Georgia, labeled “ GEORGIA: Walker Co. / 9 km W Villanow, Hwy 136 / Taylor Ridge, 640m, at lights / 4-V-1996, D. Yanega [printed, white paper] // [circular blue disc signifying dissection by RJB] // INHS / Insect Collection / 408,101 [printed, white paper with black border] // NEOTYPE / Pachybrachys / subfasciatus LeConte 1824 / design. R. J. Barney, 2016 [printed, red paper]”, is herein designated as the neotype and deposited in INHS. The specimen is point-mounted and dissected, with the abdomen and genitalia affixed to the point. It is in excellent condition, with all appendages intact ( Fig. 1).

Suffrian (1852) attributes authorship of P. biguttatus to Norwich in Sturms Catalog. Fall (1915) made no mention of P. biguttatus , and no other information is available.

Fall (1915) reported examining a female type (holotype?) of P. impurus , sent to him by Professor Taschenberg, as being from Texas. Suffrian (1852) must have examined multiple specimens as he cited this species as “In Texas ( Mus . Deyrolle) and North America without specifying the discovery site ( Mus . Mannerh. communicated by Dejean as P. femoratus Oliv. ” Neither MNHUB nor MLUH are in possession of Suffrian’ s type. Therefore, a male specimen from Texas is herein designated as neotype and deposited at TAMU. It is labeled, “ TEXAS: Brazos Co. / College Station, Lick / Ck. Pk., III-11-1999 / E. G. Riley-737, UV [printed, white paper] // [circular pink disc signifying dissection by EGR] // NEOTYPE / Pachybrachys / impurus Suffrian 1852 / design. R. J. Barney, 2016 [printed, red paper]”. The specimen is pointmounted and dissected, with the abdomen and genitalia affixed to the point. It is in excellent condition, with all appendages intact ( Fig. 9A).

A female syntype of P. xanthias , labeled “24087 [printed black on white paper] // Pachybrachys / xanthias Suffr. / Missouri Koch. [hand-inked, green paper] // SYNTYPUS / Pachybrachis / xanthias Suffrian, 1852 / labelled by MNHUB 2015 [printed, red paper]”, was examined and photographed ( Fig. 9C). Suffrian (1852) reported the type localities as Pennsylvania and Missouri. However, this female specimen has distinct ocular lines, is not P. subfasciatus , and, therefore, is a name of uncertain application.

Fall’ s male holotype of P. impurus var. umbrosus , labeled, “Salina / Ks. VI // ♂ // H. C. FALL / COLLECTION [printed, white paper] // TYPE / var. / umbrosus [hand-inked, white paper] // M.C.Z. / Type [printed, red paper] / 24984 [hand-inked]”, was examined and photographed ( Fig. 9B).

Redescription. Male. Black to brown, pronotum at sides, broad, irregular, transverse median elytral fascia, and elytral apices red or yellow; L = 1.97– 2.35 mm (mean = 2.20 mm, n = 10); W = 0.98– 1.26 mm (mean = 1.15 mm, n = 10); L/W = 1.85–1.99 (mean = 1.91, n = 10). Head: Not wider than thoracic apex, HW = 0.63–0.79 mm (mean = 0.73 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.13–0.19 mm (mean = 0.16 mm, n = 10); IOD/HW = 0.19–0.25 (mean = 0.22, n = 10); face entirely black to primarily yellow with median brown macula between eyes, more punctate in darker areas ( Figs. 1, 9A–B). Pronotum: Anterior width much less than basal width; pale areas reddish yellow, may be confined to sides or greatly expanded ( Figs. 1, 9A–B); PL = 0.87–1.04 mm (mean = 1.01 mm, n = 10); PW = 0.58–0.74 mm (mean = 0.67 mm, n = 10); PL/PW = 1.40–1.62 (mean = 1.50, n = 10). Elytra: Punctures confused baso-medially, striae fragmented or irregular ( Fig. 1). Pygidium: Finely punctate, weakly convex; entirely black to brown with large, confluent, oval yellow macula occupying majority of surface. Venter: Black; upper sides of last ventrite yellow. Legs: Brown, femora basally and tibiae distally yellow; front claws distinctly enlarged. Genitalia: Median lobe in en-face view narrow and long, with a large PRL and elongated OS; DEO composed of a pointed terminus, with excurvate ALM with denticles, appearing as a median point with 2 lateral points; in lateral view, shaft very long and inflated, creating a long tube with 3-point DEO ( Figs. 1, 9A–B) (56 males from 17 states were dissected).

Female. As in male, except L = 2.40–2.70 mm (mean = 2.53 mm, n = 10); W = 1.25–1.49 mm (mean = 1.36 mm, n = 10); HW = 0.79–0.90 mm (mean = 0.85 mm, n = 10); IOD = 0.22–0.30 mm (mean = 0.25 mm, n = 10); IOD/HW = 0.26– 0.33 (mean = 0.30, n = 10).

Remarks. Pachybrachis impurus and P. impurus var. umbrosus are synonomized with P. subfasciatus . I cannot confirm P. xanthias as synonymous based upon the syntype available to me. Fall (1915) speculated that “ Impurus, xanthias , and umbrosus are almost surely forms of one variable species, but until the males are more perfectly connected I do not feel quite safe in uniting them with subfasciatus ”. The dissection of 56 specimens from 17 states, including specimens idenfied by H. C. Fall as P. subfasciatus and P. impurus , revealed minor variability, but no significant differences.

Distribution. Pachybrachis subfasciatus is most abundant in the Middle Atlantic States to Michigan and Illinois, and south to the Gulf of Mexico. The paler “ impurus ” version is scattered across the Great Plains (Map 1A). New state records were confirmed for two states: New Hampshire and Tennessee.

Biological Notes. Lovell (1915) reported P. subfasciatus taken on Prunus virginiana L. ( Rosaceae ) in Hopkinton, Massachusetts. Johnson (1915) recorded P. subfasciatus on elm ( Ulmus sp. , Ulmaceae ) in Meriden, Connecticut. In my study, label citations were found pertaining to plant associations from various states: Quercus sp. and Corylus sp. ( Betulaceae ) in Massachusetts; Crataegus sp. ( Rosaceae ) and Carya sp. ( Juglandaceae ) in Illinois; Vaccinium sp. ( Ericaceae ) in Maryland; and Diopyros virginia L. ( Ebenaceae ) in Missouri. Specimens were reportedly collected in the following traps: Leggett, blacklight and MV, canopy, boll weevil pheromone, and Malaise.

Very little is known biologically of adult P. subfasciatus , and even less is known of the immature stages ( Lawson 1976; LeSage 1985). During my study, a pair of specimens from Suffolk County, New York had pupal cases glued to the points in addition to the emerged adults ( Fig. 9D). This is the first example of the pupal case of a North American Pachybrachis that I have seen.

Specimens Examined. See Appendix 1.

2. Pachybrachis dilatatus Suffrian, 1852 ( Fig. 2, Map 1A)

Pachybrachys dilatatus Suffrian 1852: 162 . LeConte 1880: 209 (taxonomy, catalogue – in error); Blatchley 1910: 1126 (taxonomy, catalogue – in error); Fall 1915: 475 (taxonomy, catalogue); Hughes 1944: 132 (catalogue – in error).

Pachybrachis dilatatus Suffrian : Downie and Arnett 1966: 1317 (taxonomy); Kirk 1969: 89 (catalogue), 1970: 86 (catalogue); Riley et al. 2003: 157 (catalogue); Clark et al. 2004: 156 (plant associations); Ciegler 2007: 178 (taxonomy).

Type Material. Suffrian (1852) described P. dilatatus as “From the eastern states”, apparently examining several specimens. A request of MLUH for information on Suffrian’ s P. dilatatus material resulted in a photograph of a pubescent specimen from a location they could not decipher. MNHUB has no Suffrian specimens labeled as P. dilatatus . Fall (1915) reported this species as from Georgia, having examined only three specimens: the holotype (unknown gender, loaned by Dr. Taschenberg) and two females from St. Catherine’ s Island, April 19 (Hubbard & Schwarz). I herein designate a neotype, labeled “GA: Upson Co. / Sprewell Bluff area / April 24, 1967 / R. Morris // Pachybrachis / subfasciatus LeConte / Det. I.S. Askevold 1988 [printed, white paper] // [circular blue disc signifying dissection by RJB] // NEOTYPE / Pachybrachys / dilatatus Suffrian 1852 / design. R. J. Barney, 2016 [printed, red paper]” and deposited in UGCA. The specimen is point-mounted and dissected, with the abdomen and genitalia affixed to a second point. It is in excellent condition, with all appendages intact ( Fig. 2).

Redescription. Male. Black, pronotum sides, large, subquadratic spot not extending to suture on elytra, and elytral apices red; L = 2.44–2.72 mm (mean = 2.57 mm, n = 10); W = 1.31–1.45 mm (mean = 1.38 mm, n = 10); L/W = 1.73–1.98 (mean = 1.86, n = 10). Head: Not wider than thoracic apex, HW = 0.77–0.89 mm (mean = 0.84 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.17–0.23 mm (mean = 0.19 mm, n = 10); IOD/HW = 0.19–0.27 (mean = 0.23, n = 10); face entirely black ( Fig. 2). Pronotum: Anterior width much less than basal width; sides with red margin narrowed behind; punctation fine and sparse ( Fig. 2); PL = 1.21–1.32 mm (mean = 1.23 mm, n = 10); PW = 0.73–0.85 mm (mean = 0.78 mm, n = 10); PL/PW = 1.50–1.68 (mean = 1.59, n = 10). Elytra: Punctures confused baso-suturally, elsewhere arranged in impressed lines; marginal interspace punctured in basal half ( Fig. 2). Pygidium: Finely punctate, black. Venter: Black. Legs: Black, front claws distinctly enlarged. Genitalia: Median lobe in en-face view with OS much wider than PRL, with elongated, distinct median and lateral basal plates ( Fig. 2); thickened ALM rounded to a triangular terminus; in lateral view, ALM recurvate with denticles projecting basally towards much inflated PRL (18 males from six states were dissected).

Female. As in male, except L = 2.57–3.06 mm (mean = 2.78 mm, n = 10); W = 1.37–1.64 mm (mean = 1.51 mm, n = 10); HW = 0.82–1.01 mm (mean = 0.91 mm, n = 10); IOD = 0.17–0.27 mm (mean = 0.23 mm, n = 10); IOD/HW = 0.20– 0.29 (mean = 0.26, n = 10).

Remarks. Pachybrachis dilatatus is one of the rarest Pachybrachis in the eastern United States and often confused with P. subfasciatus . Fall (1915) only had access to three specimens: the Suffrian type and two females, all from Georgia. Reports of P. dilatatus in the District of Columbia, Indiana ( Blatchley 1910), New Jersey, New York ( LeConte 1880), Ohio ( Hughes 1944), and Pennsylvania cannot be confirmed, and most likely are P. subfasciatus ( Riley et al. 2003) . The two species are superficially very similar in appearance; however, the differences in size, range, and aedeagus enable their identification. Two instances of P. subfasciatus and P. dilatatus in the same collecting event were found (Alabama and Mississippi).

Distribution. Pachybrachis dilatatus is found in the southeastern coastal states from North Carolina to Alabama (Map 1A). A new state record was confirmed for Alabama.

Biological Notes. This species has been recorded from Quercus sp. ( Kirk 1969, 1970; Wilcox 1979) as reported by Clark et al. (2004). However, those identifications cannot be confirmed, and no plant associations were found on any labels in this pres- ent study. Specimens were reportedly collected in the following traps and habitats: Leggett, blacklight in mixed mesic forest, tanglefoot screen, boll weevil pheromone, and cotton DVAC sample.

Specimens Examined. See Appendix 2.

3. Pachybrachis femoratus ( Olivier, 1808) ( Figs. 3, 9E–F, Map 1B)

Cryptocephalus femoratus Olivier 1808: 810 . Cryptocephalus femoratus Say 1824: 439 (in error). Cryptocephalus sparsus Newman 1840 (1841): 79. Pachybrachys characteristicus Suffrian 1852: 176 .

Pachybrachys femoratus, var. aquilonis Fall 1915: 375 .

Pachybrachys sparsus (Newman) : Haldeman 1849: 261 (taxonomy, catalogue); Melsheimer 1853: 127 (catalogue); LeConte 1880: 209 (catalogue); Balsbaugh and Hays 1972: 31 (catalogue).

Pachybrachys femoratus (Olivier) : Haldeman 1849: 261 (taxonomy, catalogue); LeConte 1880: 208 (taxonomy, catalogue); Blatchley 1910: 1127 (taxonomy, catalogue); Fall 1915: 374 (taxonomy, catalogue); Hughes 1944: 131 (catalogue); Wilcox 1954: 390 (catalogue).

Pachybrachis characteristicus Suffrian : Fall 1915: 375 (taxonomy, catalogue); Balsbaugh and Hays 1972: 31 (catalogue); Clark et al. 2004: 156 (plant associations).

Pachybrachis femoratus (Olivier) : Melsheimer 1853: 126 (catalogue); Downie and Arnett 1966: 1319 (taxonomy); Kirk 1970: 86 (catalogue); Balsbaugh and Hays 1972: 31 (catalogue); Riley et al. 2003: 157 (catalogue); Clark et al. 2004: 156 (plant associations); Ciegler 2007: 178 (taxonomy).

Type Material. Olivier (1808) description and plate figure of P. femoratus , reported as from “ Carolina ”, is too vague to be definitive. Fall (1915) made no mention of having seen a type, and finding “Olivier’ s description of little value”, he decided, “to accept as femoratus the species so labeled in the LeConte Collection”. Therefore, I herein designate a neotype, labeled “ DELAWARE: Sussex / Co., Lewes / 12 June 1983 / G.F.&J.F. Hevel [printed, white paper] // [circular blue disc signifying dissection by RJB] // NEOTYPE / Cryptocephalus / femoratus Olivier 1808 / design. R. J. Barney, 2016 [printed, red paper]” and deposited in MCZ. The specimen is pointmounted and dissected, with the abdomen and genitalia affixed to a second point. It is in excellent condition, with all appendages intact ( Fig. 3). There are additional specimens (6♂♂ 3♀♀) with identical collecting data.

Newman (1840) described C. sparsus as “ black, obliquely speckled white, head and prothorax punctate, elytra profoundly punctate, punctures arranged”. The specimen was taken by a Mr. Doubleday at Trenton Falls, Oneida County, New York and housed in the cabinet of The Entomological Club of London. The first reference to P. sparsus as a synonym of P. femoratus that I have found was in Balsbaugh and Hays (1972). However, I have never encountered an author make mention of examining the holotype, and the description applies to several species of Pachybrachis .

Two male syntypes of P. characteristicus , labeled “24077 [printed, white paper] // Hist.-Coll. ( Coleoptera ) / Nr. 24077 / Pachybrachys / characteristicus Suffr. / Americ. Sept. Knoch. / Zool. Mus . Berlin [printed, green paper with black line border] // SYNTYPUS / Pachybrachis / characteristicus Suffrian, 1852 / labelled by MNHUB 2015 [printed, red paper]”, and one female syntype labeled “24077 [printed, white paper] // characteristi / cus N / Suffr. / Missouri Kn. [hand-written, green paper with black line border] // SYNTYPUS / Pachybrachis / characteristicus Suffrian, 1852 / labelled by MNHUB 2015 [printed, red paper]” were examined. I herein designate one of the male syntypes mentioned above as a lectotype, with the additional label, “ LECTOTYPE / Pachybrachys / characteristicus / Suffrian 1852 / design. R. J. Barney, 2016 [printed, black on red paper]” ( Fig. 9F).

Fall’ s male holotype of P. femoratus var. aquilonis , labeled “ ♂ // C.A. Frost / Framingham / VI 2 -06 Mass. // H. C. FALL / COLLECTION [printed, white paper] // TYPE / var. / aquilonis [hand-inked, white paper] // M.C.Z. / Type [printed, red paper] / 24915 [hand-inked]”, was examined and photographed ( Fig. 9E).

Redescription. Male. Small to medium size; mottled brown and yellow; L = 1.90–2.18 mm (mean = 2.05 mm, n = 10); W = 1.03–1.18 mm (mean = 1.12 mm, n = 10); L/W = 1.81– 1.87 (mean = 1.83, n = 10). Head: Not wider than thoracic apex, HW = 0.63–0.73 mm (mean = 0.70 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.10–0.19 mm (mean = 0.16 mm, n = 10); IOD/HW = 0.15–0.26 (mean = 0.22, n = 10); face mostly yellow with brown to black, often discontinuous, W-shaped macula connecting the vertex and base of antennae, and up to base of upper lobe of eyes; antennae brown, not reaching elytral declivity ( Fig. 3). Pronotum: Anterior width much less than basal width; yellow with brown markings broad and diffuse, pale areas small, side margins not smooth or partially so ( Fig. 3); PL = 0.89–1.03 mm (mean = 0.96 mm, n = 10); PW = 0.60–0.72 mm (mean = 0.67 mm, n = 10); PL/PW = 1.39–1.55 (mean = 1.45, n = 10). Elytra: Yellow with brown maculae in equal proportion to yellow in no particular pattern; punctation confused in baso-sutural region, striae weak and incomplete basally, mostly regular laterally and apically ( Fig. 3). Pygidium: Finely punctate, weakly convex; brown with large, confluent, oval yellow macula occupying majority of surface. Venter: Brown, upper sides of last ventrite yellow. Legs: Brown, femora yellow basally, tibiae distally; front claws distinctly enlarged. Genitalia: Median lobe in en-face view compressed with circular OS and darkly sclerotized median basal plate; en-face outline with ALM broadly and evenly rounded to a thick triangular nodule. In lateral view, ventral surface broadly inflated, distinctly narrowed at basal contraction; beard consisting of long setae at ALM ( Fig. 3) (92 males from 21 states were dissected).

Female. As in male, except L = 2.26–2.55 mm (mean = 2.35 mm, n = 10); W = 1.26–1.47 mm (mean = 1.34 mm, n = 10); HW = 0.77–0.89 mm (mean = 0.82 mm, n = 10); IOD = 0.14–0.25 mm (mean = 0.23 mm, n = 10); IOD/HW = 0.19–0.31 (mean = 0.27, n = 10).

Remarks. Fall (1915) recognized P. aquilonis as a darker variety of P. femoratus in the northeastern United States and P. characteristicus as a lighter species in Florida, which “may yet be necessary to reduce... to varietal standing”. A comparison of aedeagal characters of 92 males from 21 states, including specimens idenfied by H. C. Fall as P. femoratus , P. characteristicus , and P. aquilonis and specimens identified by E. G. Riley as EGR#74, EGR#94, and EGR#98, revealed minor variability but no significant differences. Therefore, P. characteristicus and P. aquilonis are synonomized as P. femoratus . Three additional potential species, temporarily categorized by E. G. Riley as EGR#74, EGR#94, and EGR#98, all from Florida, are now recognized as P. femoratus . Pachybrachys sparsus is a name of uncertain application.

Distribution. Primarily found in the coastal states from Mississippi through Florida, north to New England, with occasional specimens found in the Midwest (Map 1B). New state records were confirmed for 12 states: Arkansas, Delaware, Illinois, Kentucky, Michigan, Mississippi, Missouri, New Hampshire, Oklahoma, Tennessee, West Virginia, and Wisconsin.

Biological Notes. There are multiple label citations in various states to Quercus spp. as a host plant for P. femoratus , which I can personally confirm in Florida and Massachusetts, including Q. alba L. (Illinois), white oak; Q. laevis Walter (Florida), turkey oak; Q. laurifolia Michx. (Florida), laurel oak; Q. nigra L. (Alabama and Florida), water oak; and Q. velutina Lam. (Massachusetts), black oak. Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg. (Florida), sand pine ( Pinaceae ), was also listed once. Clark et al. (2004) reported several additional published potential plant associations in the United States, and even Mexico, but the Pachybrachis species cannot be confirmed. The following habitats and traps were cited on labels: xeric oak scrub, rosemary sandhill, longleaf pine/ turkey oak/wiregrass, sand pine/rosemary scrub, Leggett trap, Malaise trap, flight trap, night sweeping, and yellow pan trap. Specimens were often collected in long series.

Specimens Examined. See Appendix 3.

4. Pachybrachis shirleyae Barney , new species ( Fig. 4, Map 1B)

Holotype. Male , labeled “TENN: Rutherford Co. / Flat Rock Cedar Glades SNA / 35.8585°N − 86.2956°W / 4.vi.2014, R. J. Barney [printed, white paper] // [circular blue disc signifying dissection by RJB] // HOLOTYPE / Pachybrachis / shirleyae / R. J. Barney, 2016 [printed, red paper]” and deposited in MCZ. The specimen is point-mounted with genitalia affixed to a second point. It is in excellent condition with all appendages intact. There are four paratypes (1♂ 3♀♀) with identical collecting data.

Paratypes. 8♂♂ 8♀♀. See Appendix 4.

Description. Male. L = 2.10–2.45 mm (mean = 2.25 mm, n = 8); W = 1.07–1.25 mm (mean = 1.16 mm, n = 8); L/W = 1.83–2.03 (mean = 1.94, n = 8). Head: Slightly wider than thoracic apex, HW = 0.79–0.88 mm (mean = 0.85 mm, n = 8); eyes bulging laterally, upper lobes widely separated, no ocular lines, IOD = 0.24–0.35 mm (mean = 0.31 mm, n = 8); IOD/HW = 0.31–0.41 (mean = 0.37, n = 8); face yellow with brown median line extending down from vertex between eyes, punctation sparse; antennae yellow and brown, not reaching elytral declivity ( Fig. 4). Pronotum: Yellow with brown to black, M-shaped macula broad, not sharply defined, median and lateral arms of macula narrowly separated from pronotal base; punctation sparse; anterior width much less than basal width, lateral margin arcuate basally ( Fig. 4); PL = 0.94–1.08 mm (mean = 1.01 mm, n = 8); PW = 0.62–0.77 mm (mean = 0.71 mm, n = 8); PL/PW = 1.35–1.58 (mean = 1.43, n = 8). Elytra: Yellow with brown to black maculae in equal proportion to yellow in no particular pattern; punctation confused in baso-sutural region, striae weak and incomplete basally, mostly regular laterally and apically ( Fig. 4). Pygidium: Finely punctate, weakly convex; brown to black with large, confluent, oval, yellow maculae occupying majority of surface. Venter: Brown to black, upper sides of last ventrite yellow. Legs: Yellow with brown femoral and tibial spots; front claws distinctly enlarged. Genitalia: Median lobe in en-face view with round to oval OS, extending apically as a groove between thickened ALM and forming a triangular extension of DEO. In lateral view, shaft very long and inflated, creating a long tube with paired beards at widest point of OS and before triangular extension of DEO ( Fig. 4) (seven males from four states were dissected).

Female. As in male, except L = 2.46–2.79 mm (mean = 2.65 mm, n = 7); W = 1.30–1.48 mm (mean = 1.41 mm, n = 7); HW = 0.92–1.06 mm (mean = 1.01 mm, n = 7); IOD = 0.41–0.52 mm (mean = 0.47 mm, n = 7); IOD/HW = 0.41–0.50 (mean = 0.46, n = 7).

Remarks. This is one of the rarer species of Pachybrachis in the eastern United States. Only 16 specimens have been examined from four states. The enlarged foreclaws, no ocular lines, and triangular extension of the DEO are a distinctive combination of characters.

Etymology. This species is named in honor and posthumous recognition of Shirley Jean Gibson Barney, MSc.

Distribution. Pachybrachis shirleyae is only known from four states: Kentucky, Mississippi, North Carolina , and Tennessee (Map 1B).

Biological Notes. The host plant is believed to be eastern red cedar, Juniperus virginica L. ( Cupressaceae ). There are multiple label citations to this effect, plus additional specimens were collected in cedar glades.

Specimens Examined. See Appendix 4.

5. Pachybrachis luridus ( Fabricius, 1798) ( Figs. 5, 10A–D, Map 2A)

Cryptocephalus luridus Fabricius 1798: 106 . Zimsen 1964: 122 (type).

Cryptocephalus femoratus Say 1824: 439 .

Cryptocephalus aesculi F. E. Melsheimer 1847: 171 .

Pachybrachys luridus, var. nigrinus Blatchley 1910: 1130 . Fall 1915: 470 (taxonomy, catalogue); Hughes 1944: 132 (catalogue).

Pachybrachys luridus (Fabricius) : Haldeman 1849: 261 (taxonomy, catalogue); LeConte 1880: 208 (taxonomy, catalogue); Blatchley 1910: 1126 (taxonomy, catalogue); Fall 1915: 469 (taxonomy, catalogue); Johnson 1915: 316 (catalogue); Dozier 1922: 118 (catalogue); Hughes 1944: 132 (catalogue); Wilcox 1954: 394 (catalogue).

Pachybrachys luridus var. festivus Fall 1915: 470 (taxonomy, catalogue); Knaus 1916 –1917: 263 (catalogue).

Pachybrachis luridus (Fabricius) : Melsheimer 1853: 126 (catalogue); Mombert 1869: 544 (catalogue); Downie and Arnett 1966: 1319 (taxonomy); Kirk 1969: 89 (catalogue); Balsbaugh and Hays 1972: 39 (catalogue); Barney 1984: 137 (catalogue); Riley et al. 2003: 159 (catalogue); Clark et al. 2004: 158 (plant associations); Ciegler 2007: 179 (taxonomy); Barney et al. 2011: 6 (catalogue), 2013: 119 (catalogue).

Type Material. Fabricius (1798) gave the source for his species as “America boreali Dom. Mauerhoff” or North America. Fall (1915) made no mention of Fabrician material, but Zimsen (1964) documented three specimens of P. luridus . An examination of these specimens ( NHMD) revealed a female P. luridus with a missing elytron ( Fig. 10A), herein designated as the lectotype and with the additional label “ LECTOTYPE / Cryptocephalus / luridus Fabricius, 1798 / design. R. J. Barney, 2016 [printed, red paper]”, a P. luridus missing most of the abdomen, and a male P. confusus .

Say (1824) gave the source for his species as “Arkansa”. Fall (1915) made no mention of Say’ s material.

Melsheimer (1847) gave the source for P. aesculi as Pennsylvania. Fall (1915) made no mention of Melsheimer’ s material. MCZ has no specimen of P. aesculi designated as a type. However, MCZ- MELS has one male specimen labeled “Melsh. [printed, white paper] / aesculi [hand-inked, white paper]”. Therefore, this specimen is designated as the lectotype ( Fig. 10B), with the additional label “ LECTOTYPE / Cryptocephalus / aesculi Melsheimer 1847 / design. R. J. Barney, 2016 [printed, red paper]”. There are four additional P. luridus specimens (4♀♀): two labeled “[circular

Map 2. Distribution of Pachybrachis species based upon specimens examined. A) P. luridus (black) and

P. turbidus (red), B) P. confusus (black) and P. sherriae (red).

blue disc] // Ziegler [printed, white paper] // lurid-us F. / Pa. [hand-inked, white paper with lined border]”; one labeled “ Pennsylvania [hand-inked, white paper] / Melsheimer [printed, white paper]”; and one labeled “Melsh. [printed, white paper] / luridus [hand-inked, white paper]”.

Blatchley’ s female holotype of P. luridus var. nigrinus , created to “distinguish them from the other black species [ Pachybrachis ] morosus and [ Pachybrachis ] carbonarius ”, is labeled “Vigo Co. / Ind. W.S.B. [printed] / 6-25-92 [hand-inked, white paper] // Purdue / Blatchley / collection [printed, white paper] // TYPE [printed, red paper] // PERC / 0084352 [printed, green paper]” ( Fig. 10D). I believe this specimen is in actuality an entirely black example of Pachybrachis nigricornis (Say) (formerly var. carbonarius ).

Fall’ s female holotype of P. luridus var. festivus , labeled “Medora, Ks. // W. Knaus. [printed, white paper] // ♀ // TYPE / festivus. [hand-inked, white paper] // M.C.Z. / Type [printed, red paper] / 24935 [hand-inked, white paper] // H. C. FALL / COLLECTION [printed, white paper]”, was examined and photographed ( Fig. 10C). Fall (1915) established P. luridus var. festivus for the primarily yellow forms found in many western states, while acknowledging that “intermediates are not lacking”.

Redescription. Male. Densely coarsely punctate, elytra mottled with yellow, especially on the sides, pronotum with anterior median line and sides red or reddish yellow, elytra varying from entirely yellow (former Pachybrachis luridus var. festivus ) to entirely black (former Pachybrachis luridus var. nigrinus ) ( Figs. 5A–B, 10A–D); L = 2.58–2.86 mm (mean = 2.68 mm, n = 10); W = 1.29–1.54 mm (mean = 1.44 mm, n = 10); L/W = 1.82–1.94 (mean = 1.86, n = 10). Head: Not wider than thoracic apex; HW = 0.79–0.92 mm (mean = 0.87 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.18–0.23 mm (mean = 0.21 mm, n = 10); IOD/HW = 0.21–0.27 (mean = 0.24, n = 10); face varying from entirely black to yellow maculate; antennae almost reaching elytral declivity ( Figs. 5, 10A–D). Pronotum: Strongly arcuately narrowed anteriorly; PL = 1.22–1.42 mm (mean = 1.28 mm, n = 10); PW = 0.81–0.95 mm (mean = 0.88 mm, n = 10); PL/PW = 1.38–1.52 (mean = 1.45, n = 10). Elytra: Coarsely punctate ( Figs. 5A–B, 10A–D). Pygidium: Black. Venter: Black. Legs: Black, front femora with oval yellow spot; front claws distinctly enlarged. Genitalia: Median lobe in en-face view with shaft widest subapically, ALM broadly rounded to terminus; basal plates weakly pigmented. Beard conspicuous, consisting of apicolateral wisp on each side; setae of wisp moderately dense, long, curved mesad. Ventral surface in lateral view inflated, without keel, with a continuous bulge from base to near terminus ( Fig. 5D) (15 males from eight states were dissected).

Female. As in male, except L = 2.79–3.25 mm (mean = 3.00 mm, n = 10); W = 1.52–1.81 mm (mean = 1.67 mm, n = 10); HW = 0.93–1.08 mm (mean = 1.00 mm, n = 10); IOD = 0.29–0.40 mm (mean = 0.33 mm, n = 10); IOD/HW = 0.29–0.37 (mean = 0.33, n = 10).

Remarks. Pachybrachis luridus was the first species of Pachybrachis in North America to receive a name. It is one of the more common and variable species, often collected in relatively long series. The pronotal and elytral coloration is highly variable. However, a consistent characteristic is the smooth anterior median line and red to reddish yellow margins of the pronotum. While there are several examples of Pachybrachis species trending from being darker in the east and paler in the west, this is not consistent with P. luridus , as considerable variability can be found within a series. Males do tend to be paler than females.

Pachybrachys moerens Stål (1857) was listed as a synonym by Balsbaugh and Hays (1972) and Riley et al. (2003). Stål gave the source for his species as Brasilia, presumably the current capital of Brazil. This species is most likely not related to P. luridus and should not be considered a synonym.

Distribution. Pachybrachis luridus can be found from New Hampshire to Florida, west to the 100 th meridian (Map 2A). New state records were confirmed for five states: Kentucky, Minnesota, Tennessee, West Virginia, and Wisconsin.

Biological Notes. A large number of plant family associations were detailed on labels, including Anacardiaceae ( Rhus glabra L.), Fabaceae ( Amorpha canescens Pursh , Baptisia tinctoria (L.) R. Br., Baptisia sp. ), Fagaceae ( Q. nigra , Quercus stellata Wangenh. , Q. velutina , Quercus spp. ), Rhamnaceae ( Ceanothus americanus L.), and Scrophulariaceae ( Penstemon cobea Nutt. ). Clark et al. (2004) reported many additional potential plant associations.

Specimens Examined. See Appendix 5.

6. Pachybrachis turbidus J. L. LeConte, 1880 ( Figs. 6, 10F, Map 2A)

Pachybrachys turbidus J. L. LeConte 1880: 208 . Fall 1915: 383 (taxonomy, catalogue).

Pachybrachis turbidus J. L. LeConte : Barney 1984: 144 (catalogue); Riley et al. 2003: 164 (catalogue); Clark et al. 2004: 162 (plant associations).

Type Material. LeConte’ s female holotype of Pachybrachis turbidus , labeled “Tex. [printed, white paper] // 403 [hand-inked, white paper] // Type [printed, white paper] / 4297 [hand-inked, red paper] // P. turbidus / Lec. [hand-inked, white paper]”, was examined and photographed ( Fig. 10F).

Redescription. Male. Medium to large, yellow with red-brown to black maculae; L = 2.74–3.23 mm (mean = 3.01 mm, n = 10); W = 1.50–1.86 mm (mean = 1.67 mm, n = 10); L/W = 1.73–1.87 (mean = 1.80, n = 10). Head: Not wider than thoracic apex; HW = 0.90–1.08 mm (mean = 1.01 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.13–0.23 mm (mean = 0.18 mm, n = 10); IOD/HW = 0.13–0.22 (mean = 0.18, n = 10); face yellow with variable reddish markings. Pronotum: Strongly arcuate, narrowed anteriorly, extensive dark red punctation covering entire area except for apical margin, narrow median anterior stripe, and margins yellow ( Fig. 6); PL = 1.40–1.66 mm (mean = 1.52 mm, n = 10); PW = 0.92–1.19 mm (mean = 1.08 mm, n = 10); PL/PW = 1.35–1.54 (mean = 1.41, n = 10). Elytra: Coarsely punctate, striae deeply impressed, marginal interspace impunctate; anterior and posterior maculae usually coalesce to form 2 irregular transverse fasciae ( Fig. 6). Pygidium: Finely punctate, weakly convex; red-brown and yellow. Venter: Reddish brown. Legs: Reddish brown; front claws distinctly enlarged. Genitalia: Median lobe in en-face view narrow and long, with a large PRL and small, oval OS; DEO culminates in a triangular extension of terminus; in lateral view, ventral surface broadly inflated, distinctly narrowed at basal contstriction; beard consisting of long setae at ALM ( Fig. 6) (10 males from four states were dissected).

Female. As in male, except L = 3.17–3.80 mm (mean = 3.52 mm, n = 10); W = 1.88–2.11 mm (mean = 2.01 mm, n = 10); HW = 1.08–1.29 mm (mean = 1.20 mm, n = 10); IOD = 0.29–0.39 mm (mean = 0.34 mm, n = 10); IOD/HW = 0.26–0.31 (mean = 0.29, n = 10).

Remarks. Pachybrachis turbidus is a robust species, primarily found on Quercus spp.

Distribution. Pachybrachis turbidus is found in the middle of the country from Texas to Nebraska. The eastern most population was discovered at Fults Hill Prairie Nature Preserve, Illinois, overlooking the Mississippi River (Map 2A). New state records were confirmed for two states: Arkansas and Nebraska.

Biological Notes. Many labels reported P.turbidus as found on Quercus spp. : Q. alba (Arkansas); Q. buckleyi Nixon & Dorr (Texas); Q. macrocarpa Michx. (Missouri); Q. texana Buckley (Illinois); Q. virginiana Mill. (Texas); and Quercus sp. (Texas). Clark et al. (2004) reported collections on Quercus fusiformis Small in Texas. Juglans microcarpa Nerl. ( Juglandaceae ), Capsicum annuum L. ( Solanaceae ), Larrea tridentata (DC.) Coville ( Zygophyllaceae ), and Zanthoxylum hirsutum Buckley ( Rutaceae ) were also label-cited for Texas.

Specimens Examined. See Appendix 6.

7. Pachybrachis confusus Bowditch, 1909 ( Figs. 7, 10E, Map 2B)

Pachybrachys proximus Bowditch 1909a: 360 (preoccupied).

Pachybrachys confusus Bowditch 1909b: 365 . Blatchley 1910: 1127 (taxonomy, catalogue); Fall 1915: 468 (taxonomy, catalogue); Hughes 1944: 132 (catalogue); Wilcox 1954: 394 (catalogue).

Pachybrachis confusus Bowditch : Downie and Arnett 1966: 1319 (taxonomy); Kirk 1969: 89 (catalogue); Balsbaugh and Hays 1972: 39 (catalogue); Riley et al. 2003: 156 (catalogue); Ciegler 2007: 178 (taxonomy); Barney et al. 2011: 6 (catalogue).

Type Material. Bowditch’ s male holotype of P. confusus , labeled “ ♂ // Natchez, Miss / June 6’95 / H. E. Weed [printed, white paper] // proximus / Type ♂ [hand-inked, white paper] // TYPE / F. C. B. Coll. [printed, pink paper] // Type [printed, white paper] / 8777 [hand-inked, red paper]”, and type female, labeled “ ♀ // Falls Church / Va. [printed, white paper] 6 – 11 [handinked, white paper] // proximus / Type ♀ [handinked, white paper] // TYPE / F. C. B. Coll. [printed, pink paper] // MCZ TYPE [printed, white paper] / 8777 [hand-inked, red paper]”, were examined and photographed ( Fig. 10E). Bowditch (1909b) realized that he had described two species of Pachybrachis under the name of P. proximus ( Bowditch 1909a) and then used P. confusus (how appropriate) as a replacement name.

Redescription. Male. Medium size to large, black with whitish yellow markings; L = 2.29– 2.47 mm (mean = 2.38 mm, n = 10); W = 1.21– 1.37 mm (mean = 1.28 mm, n = 10); L/W = 1.80–1.92 (mean = 1.86, n = 10). Head: Slightly wider than thoracic apex; HW = 0.80–0.93 mm (mean = 0.87 mm, n = 10); eyes widely separated, no ocular lines, IOD = 0.32–0.43 mm (mean = 0.36 mm, n = 10); IOD/HW = 0.37–0.49 (mean = 0.42, n = 10); face yellow with black median line extending down from vertex between eyes, forming distinctive paired, yellow inverted triangular maculae; punctation sparse; antennae black, not reaching elytral declivity face ( Fig. 7). Pronotum: Yellow, with extensive black punctation covering entire area except for apical margin, narrow median anterior stripe, and possible marginal spots yellow ( Fig. 7); PL = 1.03–1.20 mm (mean = 1.11 mm, n = 10); PW = 0.78–0.90 mm (mean = 0.84 mm, n = 10); PL/PW = 1.25– 1.36 (mean = 1.33, n = 10). Elytra: Black, basal margin raised with adjacent areas, a dot-like spot at mid-elytra and apex, yellow; punctation confused and coarser than prothorax, possible fragments of lateral striae ( Fig. 7). Pygidium: Finely punctate, weakly convex; black, sometimes with small, paired, oval. yellow maculae. Venter: Black, sometimes upper sides of last ventrite yellow. Legs: Black, femora yellow basally, tibiae distally; front claws distinctly enlarged. Genitalia: Median lobe in en-face view with ALM gently curved to rounded terminus; OS circular, basal plates sclero- tized; beard with long setae visible from ALM to terminus; in lateral view, shaft inflated but truncated ( Fig. 7) (27 males from seven states were dissected).

Female. As in male, except L = 2.59–2.84 mm (mean = 2.70 mm, n = 10); W = 1.39–1.59 mm (mean = 1.48 mm, n = 10); HW = 0.94–1.04 mm (mean = 0.99 mm, n = 10); IOD = 0.41–0.54 mm (mean = 0.47 mm, n = 10); IOD/HW = 0.42–0.52 (mean = 0.47, n = 10).

Remarks. Pachybrachis confusus is a relatively rare species in the southeastern United States. Superficially, it could be confused with Pachybrachis stygicus Fall or Pachybrachis atomarius (F. E. Melsheimer) , but P. confusus is more robust, has enlarged foreclaws, paired pale, inverted triangular areas between the eyes, and a distinctive aedeagus.

Distribution. Specimens of P. confusus from Maryland to Mississippi were examined. New state records were confirmed for seven states: Arkansas, Florida, Kentucky, Maryland, Massachusetts, Tennessee, and West Virginia (Map 2B).

Biological Notes. A series (5♂♂ 5♀♀) of P. confusus was labeled as collected from Desmodium sp. ( Fabaceae ) on a pine forest roadside in South Carolina .

Specimens Examined. See Appendix 7.

8. Pachybrachis sherriae Barney , new species ( Fig. 8, Map 2B)

Holotype. Male , labeled “ USA: IL: Monroe Co. / Fults Hill Prairie Nat Pre / 38.1544°N 90.1848°W / 1.vii.2013 R. J. & S. Barney [printed, white paper] // [circular blue disc signifying dissection by RJB] // HOLOTYPE / Pachybrachis / sherriae / R. J. Barney, 2016 [printed, red paper]” and deposited in MCZ. The specimen is point-mounted with genitalia affixed to a second point. It is in excellent condition with all appendages intact. There are 23 paratypes (5♂♂ 18♀♀) with identical collecting data.

Paratypes. 188♂♂ 193♀♀. See Appendix 8.

Description. Male. L = 2.03–2.28 mm (mean = 2.15 mm, n = 10); W = 1.07–1.22 mm (mean = 1.14 mm, n = 10); L/W = 1.79–1.97 (mean = 1.89, n = 10). Head: Slightly wider than thoracic apex; HW = 0.71–0.79 mm (mean = 0.74 mm, n = 10); eyes not widely separated, no ocular lines, IOD = 0.11–0.21 mm (mean = 0.16 mm, n = 10); IOD/HW = 0.16–0.27 (mean = 0.22, n = 10); face yellow with brown median line extending down from vertex between eyes, punctation sparse; antennae yellow and brown, not reaching elytral declivity ( Fig. 8). Pronotum: Yellow with reddish brown to black, M-shaped macula broad, not sharply defined, median and lateral arms narrowly separated from pronotal base; punctation sparse; anterior width much less than basal width, lateral margin arcuate basally; PL = 0.96–1.07 mm (mean = 1.02 mm, n = 10); PW = 0.63–0.75 mm (mean = 0.68 mm, n = 10); PL/PW = 1.40–1.54 (mean = 1.49, n = 10). Elytra: Yellow with brown to black maculae in equal proportion to yellow in no particular pattern; punctation confused in baso-sutural region, striae weak and incomplete basally, mostly regular laterally and apically ( Fig. 8). Pygidium: Finely punctate, weakly convex; brown to black with large, confluent, oval yellow macula occupying majority of surface. Venter: Brown to black, upper sides of last ventrite yellow. Legs: Yellow with brown femoral and tibial spots; front claws distinctly enlarged. Genitalia: Median lobe in en-face view near equal width throughout, ALM gently curved to terminus; OS ca. 2 times longer than wide, median basal plate pigmented; PRL very small; beard inconspicuous with tips of longest setae visible in en-face view, more obvious in lateral view as apicolateral space patch, setae long, straight. Ventral surface inflated but narrowing quickly to terminus due to short PRL ( Fig. 8) (33 males from seven states were dissected).

Female. As in male, except L = 2.14–2.53 mm (mean = 2.37 mm, n = 10); W = 1.10–1.35 mm (mean = 1.27 mm, n = 10); HW = 0.75–0.87 mm (mean = 0.83 mm, n = 10); IOD = 0.19–0.26 mm (mean = 0.23 mm, n = 10); IOD/HW = 0.22–0.33 (mean = 0.28, n = 10).

Etymology. This species is named for the cocollector of the holotype and a long-time advisor and inspiration to the author, Sherri Ann Zellers Barney, RN.

Remarks. Pachybrachis sherriae resembles many other species in Fall’ s (1915) Group C, which are yellow and confusedly mottled with brown or black markings. However, the enlarged foreclaws, lack of ocular lines, and distinctive aedeagus permit identification. Many specimens were temporarily categorized by Edward G. Riley (EGRC) as EGR#31.

Distribution. Pachybrachis sherriae is known from Illinois to Nebraska, south into Texas (Map 2B).

Biological Notes. Large series of specimens were collected from Desmanthes sp. ( Fabaceae ) in Kansas and Lousiana and Senegalia (Acacia)

22 THE COLEOPTERISTS BULLETIN 71(1), 2017

greggii (A. Gray) Britton & Rose ( Fabaceae ) in Texas. A single female was reported from R. glabra in Illinois.

Specimens Examined. See Appendix 8.