Hemicrepidius melanopthalmus (Melsheimer, 1846)

Etzler, Frank E., 2020, A Revision of the Genus Hemicrepidius Germar, 1839 (Coleoptera: Elateridae) of the New World, with Comments on Global Classification, The Coleopterists Bulletin (mo 18) 74, pp. 1-126 : 47-50

Hemicrepidius melanopthalmus (Melsheimer, 1846) , status resurrected

( Figs. 22a–e , 23b )

Athous melanopthalmus Melsheimer 1846: 154 .

Asaphes melanophthalmus (Melsheimer) : LeConte 1853: 450; Candèze 1863: 221; Horn 1880: 75 (as synonym of H. bilobatus ). [unjustified emendation]

Athous cavifrons Melsheimer 1846: 154 .

Asaphes cavifrons (Melsheimer) : LeConte 1853:453; Candèze 1863: 223 [not examined by Candèze]; Horn 1880: 75 (as synonym of H. bilobatus ).

Asaphes tener LeConte 1853: 452 ; Candèze 1863: 221 (as synonym of H. melanopthalmus ); Horn 1880: 75 (as synonym of H. bilobatus ).

Diagnosis. Hemicrepidius melanopthalmus can be distinguished from all other Hemicrepidius species by the lack of a carina on the pronotal hind angle, the lack of sublateral basal notches, a sinuate posterior margin of the hypomeron, and an aedeagus with a lateral compression of the parameres basally to the pre-apical expansion.

It is most similar to H. bilobatus , and the ways to distinguish the two species are provided in the Diagnosis for that species above.

Redescription. Length 12.0–19.0 mm, width 3.0–5.0 mm; color uniformly light red-brown to dark red-brown, elytra not contrasting with pronotum, legs and antennae paler in most, almost yellow-brown, paler on ventral side; integument shining. Setae short, mostly depressed, a few setae erect, yellow. Head: Punctures subumbilicate, nearly contiguous; frons weakly concave anteromedially, some with slight triangular impression medially in dorsal view; frontal carina roughly Vshaped, sometimes interrupted medially by punctures when viewed from front, frontoclypeal region absent medially, at most weakly projecting when viewed from side; eyes large and strongly bulging, ocular index 58–65 (n = 4); antenna weakly serrate, antennomere 2 subcylindrical, antennomere 3 subcylindrical, 1.5× length of 2 and with similar punctation, antennomere 3 subequal in length to 4, antennomeres 4–11 about 3× longer than wide, each with medial line bearing less setae than edges, antennomere 11 oval or with tip slightly constricted, antenna reaching pronotal posterior edge to passing it by 2 antennomeres. Prothorax: Pronotum as wide as long to 1.2× wider than long, widest at hind angles (most males) or just anterior of midlength (most females); sides almost subparallel laterally and narrowing at anterior one-fourth in males, broadly arcuate in females; hind angles usually subparallel, sometimes appearing slightly divergent due to sinuate edge anterior of hind angle, dorsal carina absent; punctures subumbilicate, similar in size to punctures on head, separated by less than puncture diameter to nearly touching on disc, punctures larger and nearly contiguous laterally; posterior edge without sublateral notches. Hypomeron with subumbilicate punctures, punctures same size as lateral pronotal punctures, nearly contiguous anteriorly and becoming less dense posteriorly; posterior margin sinuate ( Fig. 22c ) with hind angle visible in lateral and ventral view at truncate tip. Prosternum with punctation slightly smaller than punctation on hypomeron, nearly contiguous laterally, less dense medially; anterior lobe weakly deflexed, broadly rounded and covering ventral mouthparts; prosternal process curved dorsad between procoxae, narrow and straight past procoxae. Pterothorax: Mesoventral cavity with lateral edges sinuate, not visible anteriad mesocoxae in lateral view; posterior edge of cavity narrowly truncate. Metaventrite with punctures smaller than punctures on hypomeron, appearing simple, nearly touching throughout. Scutellar shield elongate, about twice as long as wide, narrowly rounded posteriorly, sides sinuate, dorsal surface weakly convex, anterior margin nearly straight, setae radiating from midline, similar in color and length to setae on pronotum and elytra or denser. Elytral striae weakly impressed basally; interstriae flat to slightly convex, punctures small but giving wrinkled appearance; basal carina simple. Legs with tarsomeres 2 and 3 moderately wide, visible dorsally, lobe on tarsomere 3 largest, lobe on tarsomere 2 slightly smaller than lobe on 3, lobe on tarsomere 4 minute, visible only ventrally, lobe on tarsomere 1 smallest, barely visible ventrally. Abdominal ventrites: Punctures simple, slightly smaller than punctures on metaventrite, separated by one puncture diameter, further apart medially; fifth ventrite broadly rounded; sides of each ventrite with microserrations. Male genitalia ( Figs. 22d, e ): Parameres subparallel laterally on basal two-thirds, greatly narrowed and laterally compressed basally to pre-apical expansions; pre-apical expansion hook-shaped; apex of paramere pointed, weakly membranous, and setose ventral patch near pre-apical expansion, 5–8 apical dorsal setae. Median lobe narrowly triangular, constricted on apical half, basal struts straight, directed latero-basally, laterally forming about 135- degree angle. Basal piece with membranous area covering two-thirds of length, transverse medially.

Distribution. Specimens were examined from the following states: UNITED STATES: AR, CT, DE, GA, IA, IL, IN, KS, LA, MA, MD, ME, MI, MO, NC, NH, NJ, NY, OH, OK, PA, RI, SC, TN, VA, WI ( Fig. 23b ). This species also occurs in Ontario, Canada (see Notes).

Notes. Below I have designated lectotypes for Athous melanopthalmus and Athous cavifrons in the Melsheimer collection in the MCZC. Along with the type of Asaphes tener , these types should solidify the concept of this species relative to H. bilobatus .

Starting with LeConte (1853), an extra “h” was added between the “p” and “th” in the specific name for H. melanopthalmus . A return to the codecompliant ( ICZN 1999) original spelling is followed here.

Due to the historical confusion of this species with H. bilobatus , it is unclear which species the specimens in Canada under the name H. bilobatus actually represent ( Bousquet et al. 2013). Based on the DNA data on the BOLD website and the DNA trees here ( Figs. 1–3 ), it appears that H. melanopthalmus is the appropriate name for many Ontario specimens, and likely for the majority of Canadian specimens previously identified as H. bilobatus .

Types Examined. Of Athous melanopthalmus : Lectotype (here designated): Female in Melsheimer collection, lacking head, ID label only; type locality Pennsylvania; in MCZC. Of Athous cavifrons : Lectotype (here designated): Male in Melsheimer collection, ID label only; type locality Pennsylvania; in MCZC. Of Asaphes tener : Holotype: Male, [pink disc]/ A. tener Lec. melanopth male?/ type 2617/ bilobata #14; type locality New York; in MCZC.

LeConte Collection (3, MCZC). [orange disc]/ bilobata #5; [pink disc]/ bilobata #6; [pink disc]/ bilobata #11.

Specimens Examined (347). UNITED STATES: Coll Janson, Ex Candèze/ ID label, Et Unis, LeConte (1, BMNH); Coll. Janson, Ex Dejean/ A. melanopthalmus var, Athous corticinus Dejean , h. in Amer. Bor., D. LeConte (3, BMNH); [green square]/ 483/ Ent Club 44-12 (1, BMNH). Arkansas: Polk Co., S of Board Camp, R29W T3S SE Sec. 22, 17-20-VIII-1985, C. B. and J. E. Barr, MV&BL (1, LSAM); Washington Co., near Fayetteville/ 20 August 1967, taken at UV light, leg. J. R. Heitzman (1, FSCA). Connecticut: [no data] (2, MCZC); New Haven Co., Mt Carmel, River Rd., 25 July 1985, leg. Joyce R. Gilbert (1, PMNH); Long Hill, VII-29-1948, VIII-3-1948, Coll. D. J. Comboni (2, PMNH); New Haven, VII-24, VIII- 4-[19]10/ J. N. Knull Collection (2, FMNH); New Haven, VIII-15-1910 (1, MCZC); Waterbry, VII-17 (1, MCZC); [New London Co.], Lyme, VII-30-1910 (1, MCZC); Windham Co., Willimantic, South Ridge Drive, Mercury vapor light, 28-August-2000, 4-August-2002, 15-August-2002, 19-20-August- 2002, leg. R. J. Pupedis (4, PMNH; DNA—M- TEC023685); Willimantic, South Ridge Drive, UV light, 4 August 2004, leg. R. J. Pupedis (1, PMNH); Hampton, at 15w BLT, August 6, 2011, leg. Eric L. Quinter/ Clarks Corner, 0.6mi ESE BM577 powerline cut, N 41°45’59.35”, W 72°0.5’6.96” (1, PMNH [ Figs. 22d, e ]). Delaware: New Castle Co., Newark Twp., University of Delaware, Ecology Woods, at light, 30 August 2013, 5 September 2013, Anthony Martin Deczynski (3, AMDC). Georgia: [no data] (3, MCZC); Union Co., Emergence Trap 20, Aug. 28, 1975, Pasture, W. W. Morill (1, MTEC). Illinois: [no data] (2, INHS830290, INHS830291); Amer. Bor, Illinois (1, BMNH); [23] (1, INHS830293); Carroll Co., 8-1-[19]51/ W. B. Price Colr (1, FSCA); [Champaign Co.], Champaign, Aug 22, 1939, Aug 1939, Sept 1, 1939, Sept 6, 1939, C. C. Mohr (5, INHS830280, INHS830281, INHS830283, INHS830501, INHS830502); Det. Blatchley, #1445 (1, INHS830502); [19569] (1, INHS830463);[18684] (2, INHS830277,INHS830478); Urbana, 10-2-1927 / frequenting oak (1, MTEC); Urbana, Sept 10, 1950, L. J. Stannard, on porch of house (1, INHS830284); Urbana, Oct 9, 1949, L. Stannard, in town (1, INHS830272); Urbana, Sept 3, 1890, Hart & Shira, at light, forest (2, INHS830273, INHS830279); [Cook Co.], Chicago, June 6 [19]08, Aug./ A. L. Melander collection (2, WSUC); LaSalle Co., 8-1935, Floyd G. Werner (1, CASC); [McHenry Co.], Algonquin, Nason (1, INHS830348); [18688] (5, INHS830276, INHS830366, INHS830368, INHS830370, INHS830474); [McLean Co.], Normal (1, INHS830269); [Peoria Co.], Peoria, Aug. 23, 1944, F. F. Hasbrouck (1, INHS830374); [Piatt Co.], Monticello, Aug. 20, 1948, Ross & Stan, Acc.No. 49514 (1, INHS 830267); Putnam Co., Aug 28, 1932, Aug. 31, 1931, May 20, 1933, M. C. Glen (3, INHS830364, INHS830413, INHS830488); Will County, Lockport, 18-7-[19]68, leg. Seaborg (1, CASC); [18685] (3, INHS830278, INHS830416, INHS830483); [7330]/[4511] (1, INHS830531); [18740] (1, INHS830482); [18745] (2, INHS830506, INHS830508); [38138] (1, INHS830510); [18744] (1, INHS830282); [10770] (1, INHS830286]; [4511] (1, INHS830285); [14021] (1, INHS830287); [7326] (1, INHS830275); [14035] (1, INHS830288); [18749] (1, INHS830289). Indiana: Allen Co., New Haven, Aug. 1985, J. P. & N. M. Downie (1, FSCA); New Haven, VIII-5-1983, VIII-5-1983, N. M. & P. J. Downie (3, FSCA); [Crawford Co.], Grantsburg, at blacklight, August 1, 1964, August 15, 1964, Dianne Eckert (2, FSCA); Howard Co., NW Howard Co, 16-VIII-1983, 17-VIII-1983, D. A. Rider/ collected at black light (2, LSAM); [Knox Co.], Bicknell, VIII-31-[19]37, E. N. Tryon (2, CASC); [Marion Co.], Indianapolis, August 3, 1962, August 1, 1963, August 14, 1964, August 16, 1965, August 27, 1969, September 1, 1969, Charles E. White (6, FSCA); Parke Co., 8-25-1961, N. M. Downie (1, FSCA); [Posey Co.], Hovey Lake, June 24, 1965, August 12, 1965, Charles E. White, Blacklight trap (2, FSCA); Tippecanoe Co., IX-6- 1970, VIII-10-1981, N. M. Downie (1, FSCA); Lafayette, 8-13-1985, 8-15-1985, 8-20-1985, 25 Aug 1990, U. V. light, N. M. Downie (4, FSCA); Wells Co., Bluffton. 23-Aug-1967 / Coll. David Wilkey (1, CASC). Iowa: [no data] (1, MCZC); [Des Moines Co.], Burlngtn (3, MCZC); Burl (1, MCZC); [Johnson Co.], Iowa City, Wickham (1, MCZC); Iowa City (1, MCZC); Iowa City, Wickham (1, CASC; 1 MCZC, MCZ-ENT00642008); Linn Co., 3.0mi N. Viola, VIII-9-12-[19]72, BL, E. L. Sleeper collr. (1, CASC); [Story Co.], 2mi S. Ames, ISU Farm, se. ¼, sec. 21, T83N, R24W, VIII- 8-12-1968, coll. pitfall trap (1, FSCA). Kansas: Jefferson Co., The Falin Property, 1.5km N jct 94 th St and Kingman Rd, 39°13’N, 95°24’W, 8-15-VIII- 2004, Z. H. Falin, ex FIT near lower meadow (2, SEMC, SM0622981, SM0622982 KUNHM-ENT); The Falin Property, 1.5km N jct 94 th St and Kingman Rd, 39°13’N, 95°24’W, 15-23-VIII-2004, Z. H. Falin, ex FIT, near upper meadow (1, SEMC, SM0625986 KUNHM-ENT); The Falin Property, 1.5km N jct 94 th St and Kingman Rd, 39°13’N, 95°24’W, 2-9-VIII-2005, Z. H. Falin, ex FIT near lower meadow, KAN1F05 171 (4, SEMC, SM0699996, SM0699999, SM0700002, SM0719097 KUNHM- ENT); The Falin Property, 1.5km N jct 94 th St and Kingman Rd, 39°13’N, 95°24’W, 9-18-VIII-2005, Z. H. Falin, ex canopy malaise, nr lower meadow, KAN1F05 184 (2, SEMC, SM0686459, SM0686460). Louisiana: E Baton Rouge Parish, Baton Rouge downtown, 14-ix-1996, H. Fei, under light (1, LSAM). Maine: [no data] (2, MCZC); York Co., West Newfield, UV Trap, 3-4 August 1983, leg. David s. MacDonald (7, PMNH). Maryland: P. G. Ritchie, 17-VII-[19]77, ex. Blacklight, C. L. Staines (1, MTEC?); Anne Arundel Co., 22-24 August 1994, C. L. Staines, blacklight (3, FSCA). Massachusetts: [no data] (2, MCZC; 2, OSUC0000551173, OSUC0000551174); E. Mass/ H. bilibatus [sic] Day (1, BMNH); [Barnstable Co.], Dennis, VII-26-[19] 56, D. Green (2, CASC); Woods Hole, 8-19-[19]10/ A. L. Melander Collection (1, WSUC); [Bristol Co.], Fall River, VIII-11-[19]07, N.S. Easton (1, MCZC); Freetown, VII-25-[19]09 (2, MCZC); [Essex Co.], Magnolia , 8-5-[18]96 (1, MCZC); [Middlesex Co.], [light brown disc = Cambridge, Mass, Hubbard and Schwarz, 1871] (1, MCZC); Concord bridge, 7-26-[19]06 (1, MCZC); Fram’- hain, VII-22-[19]06, C. A. Frost (1, FSCA; 1, MCZC); Framingham, 8-1-[19]51, C.A. Frost/ At light (1, MCZC); Melrose, Dodge Collection (1, CASC); Melrs Hghds, 8/19/[19]17/ Stockwell’ s Moth trap (1, PMNH); At Arc light, Melrose Hlds, 7-19-[19]20/ J. E. R. Holbrook collector (1, PMNH); Melrs Hghds, 7/31/[19]17/ Stockwell’ s Moth Trap (1, PMNH); Sherborn, July-23-1892, Jul-1 (2, MCZC); Tyngs, 7-17-[18]92, July-[18]97, July 1900, 11-VII, July-19’/ H.F.B. (5, MCZC); Tyngsboro 03-VIII-[19]03, Aug-[19?]03, 06-VIII-[19]28, 06- VIII-[19]29 (5, MCZC); Tyngsboro, from well, 03- VIII-30, 06-VII-30, ’06 VIII, 07-VIII-5, 08-VIII-8/ Blanchard collection (5, MCZC); Tyngs, 8-4-[18]96 (1, MCZC); Tyngsboro 8-11-[19]04, 8-8-[19]15 (3, MCZC); Tyngsboro, ‘21 Aug/ Blanchard/ A. Fenyes Collection (1, CASC); Tyngs/ Blanchard/ A. Fenyes Collection (1, CASC); Wakefield, VIII-3 (1, MCZC); [Nantucket Co.], Natuck, July-30-1894 / A.P. Morse (1, MCZC); Nantucket Isl./ [red square]/ Bolter Coll., No. E7963e (1, INHS830295); [Norfolk Co.], Brookline (4, MCZC); Cohasset, Aug-1- [19]05, Aug-15-[19]08, Aug-15, Aug-20 (5, MCZC); Milton, 7-21-[18]95 (1, MCZC); Nor. Co./ Hayward Collection (4, MCZC); [Plymouth Co.], Marshfield, VIII-5 (1, MCZC); Marion, Aug-1-[19]03, Aug-13- [19]03, 7-20 (3, MCZC); Marion, Bowditch/ A. Fenyes Collection (6, CASC); [Suffolk Co.], Dorchester, Jul-29-1905 (1, MCZC); [County?] Teurla [?], 7-28-[18]72 (1, MCZC). Michigan: Oakland Co., VIII-23-1931, A. W. Andrews (1, MCZC); Washtenaw Co., Ann Arbor, VIII-15-1924, F. M. Gaige/ taken at light (3, FSCA); Ann Arbor, at porch light, VIII-2-1927, W. W. Newcomb (1, FSCA); Ann Arbor, VII-29-1953, VIII-8-[19]53, VIII-12- [19]53, G. H. Nelson/ Collected at light (4, FSCA); Van Buren Co., Kendel, 21 May 1986, J. A. Shuev colr. (1, MTEC). Missouri: [no data] (1, INHS830271); St Louis Co., 1 mile North of Eureka on August 17 th, 1979, S. Jay Hanselmann, coll. (1, BYUC). New Hampshire: [no data] (1, MCZC); [County?], Ramper, VII-24-1925, Darlington/ flying at night (1, MCZC). New Jersey: [no data] (4, MCZC); [Essex Co.], Livingston, Sept. 2, 1950, V. Roth (1, OSAC0000566838). New York: [no data] (1, BMNH; 3, CASC; 1, WSUC); Albany Co., Rensselaerville Huyck Preserve/ Ross H. Arnett Jr. Collector, August 1, 1975 (1, FSCA); Columbia Co., nr. Kinderhook, Oxycopis Pond/ Ross H. Arnett Jr. Collector, 15-VII-1981 (1, FSCA); [Essex Co.], Ticonderoga, Aug. 6, 1929, F. C. Listman (1, PMNH); Ontario Co., nr. Victor/ See note no. 276, 1956, R. H. Arnett, Jr. (1, FSCA); [Richmond Co.], Staten Is., IX-1901 (1, CASC); Suffolk Co., Gardiner’ s Island, August 18, 2013, leg. R. J. Pupedis/ baited pitfall traps, meadow nr. Tob. Pd., N 41°5’25.83”, W 72°5’28.66” (3, PMNH, YPM ENT. No 828275, YPM ENT. No 828279, YPM ENT. No 828281; DNA—MTEC023683,MTEC023684); [Tompkins Co.], Ithaca, 17-Aug-[18]89 (1, MCZC); Ithaca, 29 July 1937, 27 July 1939, J. G. Franclemont (3, FSCA); Ithaca, 8 Aug. [18]90/ Van Dyke Collection (1, CASC); Ringwood Res., 24 July 1964 / R. Silverglied (1, MCZC). North Carolina: [Wake Co.], Apex, 26-VI-[19]49, Chermock (1, CASC). Ohio: Clinton Co., VIII-28-[19]55, IX-25- [19]55 [ Fig. 22c ]/ Collr. E. I. Hazard (2, FSCA); Franklin Co., VIII-27-[19]60/ Collr. E. I. Hazard (1, FSCA); Columbus, IX-1-[19]56, VIII-8-[19]64/ D. J. & J. N. Knull Collrs. (1, FMNH; 1, FSCA); [Hamilton Co.], Cin[cinnati], 8/17/[19]00/ Van Dyke Collection (2, CASC); [Knox Co.], Gambier, VIII-6-[19]44 (1, FSCA); [Muskingum Co.], Zanesville, 6-17-[19]18, 7-14-[19]18 [ Fig. 22a ], A. E. Miller (2, FSCA); [Wayne Co.], Wooster, 07-16- [19]20, 07-20-[19]20, R. Miller (2, FSCA). Oklahoma: Latimer Co., VIII-[19]86, IX-1986, VIII-1987, IX-1987, VII-1988, IX-1988, VIII- 1991, VIII-1992, VIII-[19]93, Karl Stephan (57, FSCA). Pennsylvania: [no data] (2, OSAC); [Berks Co.], Stony Crk Mills, VIII-10-[19]48, D. G. Kissinger (1, FSCA); [Dauphin Co.], Harrisburg (3, MCZC); Rockville, VII-30-1911, H. B. Kirk (1, MCZC); Rockville, VII-27-[19]11/ J. N. Knull Collection (1, FMNH), Rockville, VIII-21-1910 / Coll. by A. Champlain (1, CASC [ Fig. 22b ]); [Lackawanna Co.], Chinchilla, VII-26-[19]10 (1, MCZC); [Northampton Co.], Easton, J. W. Green, VII-30-[19]29 (1, CASC). Rhode Island: [Washington Co.], Watch Hill, July 22, 1909, W. Robinson (1, CASC). South Carolina: [Pickens Co.], Clemson, IX-15-[19]31/ D. Cartwright (1, FMNH); 02 Creekside Drive, N34.696889, W82.824169, September 2015, Anthony Martin Deczynski (1, AMDC). Tennessee: Clay Co., Standing Stone State Park (1, PMNH); Sevier Co., GSMNP, Twin Creeks ATBI Plot 273819E 3951793N, 31-Jul-15-Aug- 2002, B. Merritt, MT0120020815 (1, LSAM0156885). Virginia: [Chesapeake I.C.], Deep Creek, VII-17- [19]52, G. H. Nelson/ Dismal Swamp/ collected at light (3, FSCA); [56855]/ J. Smart (1, BMNH). Wisconsin: [Dane Co.], Madison, 9-15-1928 / C. E. Woodworth collector (2, WSUC); Madison, Aug 1927, leg. Henry Bunting (1, PMNH); Sauk Co., Pine Hollow S. N. A., 42°22’33”N, 89°54’11”W, 20-30 July 2000, A. S. Ramsdale Colr., F.I.T. mesic Acer forest (1, MTEC); Hemlock Draw, T10N/R5E/ Sec.07, July 26-Aug. 3, 1999, Alistair Ramsdale/ Flight Intercept Trap, mesic hardwood forest (1, MTEC); Milwaukee, 4-8-[19]35, Harry van Emden (1, BMNH). Unknown locality: 7-20-[18]84, G(?). H. (1, MCZC); Brewer, Jul-11-[18]79 (1, MCZC); ID label only (1, MCZC).

DNA Specimen. CANADA: Ontario: Thousand Islands National Park (BIOUG06825).

Figures

Fig. 22. Hemicrepidius melanopthalmus. Dorsal habitus: a) Male, b) Female. c) Pronotum, lateral view. Male genitalia: d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

Fig. 23. Distribution maps of examined Hemicrepidius specimens (blue/gray dots). a) H. indistinctus, b) H. melanopthalmus.

Figs. 1–2. 1) Bayesian tree of CO1 and 28S; values shown are posterior probabilities; Elaterinae clade not to scale, shortened to fit in figure; scale bar represents 4% genetic variance. 2) Maximum likelihood tree of CO1 and 28S; values shown are ultra-fast bootstrap; Melanotus clade not to scale, shortened to fit in figure; scale bar represents 5% genetic variance. Both figures: Branches with support values below 80 were collapsed. Red clade represents Hemicrepidius species. Blue arrows indicate Central American species removed from Hemicrepidius and placed into Athous.

Fig. 3. Maximum likelihood tree of CO1 barcode region. Values shown are ultra-fast bootstrap values. Branches with values below 80% are collapsed. Only selected basal branches and Hemicrepidius values are given. Blue clades represent Athous species. Green clades represent species in the Limonius group of genera (Etzler 2019). Purple clades represent genera in Dendrometrini (Cate 2007). Red clade represents Hemicrepidius species. Black clades are remaining groups. Blue arrows indicate Central American species removed from Hemicrepidius and placed into Athous. Scale bar represents 7% genetic variance. Taxa and groups given in Tables 1 and 2.