Hemicrepidius tumescens (LeConte, 1861)

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 : 85-120

Hemicrepidius tumescens (LeConte, 1861)

( Figs. 9b, d , 40a–e , 41b )

Asaphes tumescens LeConte 1861: 348 ; Candèze 1863: 224 [not examined by Candèze]; Horn 1880: 72.

Hemicrepidius tumescens (LeConte) : Leng 1920: 170; Van Dyke 1932: 444; Lane 1952: 66; Hatch and Kincaid 1958: 14; Bousquet 1991: 183; Bousquet et al. 2013: 184.

Diagnosis. Hemicrepidius tumescens can be distinguished from all other species of

Hemicrepidius by the posterior margin of the hypomeron emarginate and with the ventral hind angle narrow, over twice as long as broad and truncate; small tarsal lobes not visible dorsally; and moderately long, gray or yellow setae.

It is very close in appearance to H. californicus and H. mescalero , and characters to distinguish H. tumescens from these species are given in their respective diagnoses.

Redescription. Length 11.0–17.0 mm, width 3.0–5.0 mm; color red-brown to brown-black, elytra not contrasting with pronotum; legs, antennae and ventral surface often paler than dorsal side; integument shining. Setae moderately long, suberect, gray, few yellow. Head: Punctures subumbilicate, nearly contiguous; integument shining; frons flat to very weakly concave anteromedially in dorsal view; frontal carina roughly V-shaped, incomplete medially when viewed from front, frontoclypeal region reduced to thin line or absent medially; eyes small and weakly bulging, ocular index 69–72 (n = 2); antenna serrate, antennomere 2 subcylindrical, antennomere 3 subtriangular, 2× length of 2, and with setation intermediate between 2 and 4, antennomere 3 subequal in length to 4, antennomeres 4–11 gradually becoming narrower apically, 2× longer than wide basally to about 3× longer than wide apically, antennomeres 4–11 densely setose throughout, antennomere 11 with tip slightly constricted, antenna reaching midlength of pronotum (females) to extending past pronotal posterior edge by 1–2 antennomeres (males). Prothorax: Pronotum as wide as long to 1.2× wider than long, widest at hind angles or just anterior of midlength; sides weakly to broadly arcuate, almost subparallel laterally, narrowing on anterior half to one-third; hind angles divergent, lateral edge sinuate anterior of hind angle, dorsal carina present, directed anteromedially, nearly confluent with lateral carina posteriorly; punctures on disc smaller than to subequal size of punctures on head, simple to subumbilicate at high magnification (50×), nearly touching to separated by one puncture diameter, punctures closer and larger laterally, nearly contiguous; posterior edge with sublateral notches. Hypomeron with subumbilicate punctures, punctures nearly contiguous anteriorly and becoming less dense posteriorly; posterior margin emarginate with hind angle visible ventrally, well projecting, narrow, twice as long as broad, truncate at tip ( Fig. 40c ). Prosternum with punctation smaller than punctation on hypomeron, simple, punctures nearly touching laterally, much less dense medially, separated by up to two puncture diameters; anterior lobe straight to weakly deflexed, short, truncate to broadly rounded, not covering or partially covering ventral mouthparts; prosternal process straight between procoxae in lateral view, moderately thickened and straight past procoxae. Pterothorax: Mesoventral cavity with lateral edges raised in near 90-degree angle, visible anteriad mesocoxae; posterior edge of cavity narrowly truncate. Metaventrite with punctures simple to subumbilicate when viewed at high magnification (50×), smaller than punctures on hypomeron, dense throughout, nearly touching. Scutellar shield elongate, about twice as long as wide, broadly rounded posteriorly, sides weakly sinuate, dorsal surface flat to weakly convex, anterior margin nearly straight, setae radiating from midline, similar in length and color to setae on pronotum and elytra. Elytral striae weakly impressed basally; interstriae flat to slightly convex, punctures small, random, giving wrinkled appearance; basal carina weakly crenulate (intermediate between Figs. 6b and 6d ), appearing simple on small specimens. Legs with tarsomere lobes narrow, relatively small, not visible dorsally, lobe on tarsomere 3 largest, narrow, only lobe visible laterally, lobe on tarsomere 2 second largest, narrow, lobes on tarsomeres 1 and 4 minute, visible only ventrally, in many difficult to see. Abdominal ventrites: Punctures smaller than punctures on metaventrite, simple, very dense, nearly touching laterally, less dense medially; fifth ventrite narrowly rounded; side of each ventrite with microserrations. Male genitalia ( Figs. 40d, e ): Parameres subparallel on basal two-thirds, sinuate basal to pre-apical expansion, apex of paramere rounded, membranous, setose ventral patch near pre-apical expansion and 4–5 dorsal apical setae. Median lobe narrow, subparallel over most of length, narrowing at apex; basal struts mostly straight, directed latero-basally at 135-degree angle, and curved just at apex. Basal piece with majority of area membranous, broadly curved basally.

Distribution. Specimens were examined from the following provinces and states: CANADA: BC; UNITED STATES: CA, OR, WA ( Fig. 41b ).

Notes. Hemicrepidius tumescens is very similar in appearance to H. californicus and has also been collected along the Pacific coast. These two species are likely sister species, with H. tumescens more common in the north, and H. californicus more common in the south. The disjunct distribution should be explored with molecular data, but specimens from Santa Cruz Island (type locality) are indistinguishable from the more northern specimens commonly collected.

Larvae have not been associated for this species.

Types Examined. Of Asaphes tumescens : Holotype: [gold disc]/ A. tumescens Lec / Type #2611; male; type locality Santa Cruz Island, California; in MCZC. Paratype: [gold disc]; female; in MCZC.

Specimens Examined (159). CANADA: British Colombia: Queen Charlotte Islands, Graham Island, mouth of Tlell River, sea level, 17-July-

1981, stop #81-35, D. H. Kavanaugh collector (2, CASC); Queen Charlotte Islands, Graham Island Tlell, 1 km S of Tlell River Mouth, 3-5m, 17-July- 1983, Stop# 83-43, D. H. and M. D. Kavanaugh (3, CASC). UNITED STATES: California: [no data]/ Coll. Janson/ A.hirtus (female), dilaticollis Motsch (1, BMNH); Humboldt Co., 12-IV-1927 / E. R. Leach, Brit. Mus. 1927-457 (3, BMNH); 11-V- 1927, E.R. Leach (2, MCZC); Samoa, VI-21-[19] 16/ F. E. Blaisdell collector (3, CASC); Somoa [sic], 23-5/ H.S. Barber (2, MCZC); Santa Barbara Co., Santa Cruz Island, vicinity Pila Christi, 2-VIII- 1969, D. C. Rentz and D. Weisman (1, CASC). Oregon: Baker Creek, 4-26-1939, K. M. and D. M. Fender (1, OSAC0000551359); [Clatsop Co.], Cannon Beach, June-9-1927 / E. C. Van Dyke collector (2, CASC); Gearhart, June-12-1933, M. C. Lane col (1, WSUC); [Curry Co.], Gold Beach, VI-10-[19]36/ Van Dyke collection (7, CASC); Lincoln Co., VI-8-[19]41/ coll’ d by F. W. Nunenmacher (5, FMNH); VI-8-[19]41, E. R. Leach (18, CASC [ Fig. 40b ]); Burntwood [sic], VIII-6-1933 / S. C. Jones coll. (1, CASC); Waldport, June-7-1925, VI-13-[19]36 [ Fig. 40c ], VII-24-1938 / E. C. Van Dyke collector (38, CASC); Waldport, 6-6-[19]26/ J. E. Davies collector (1, OSAC0000836988); [Tillamook Co.], Pacific City, 5-16-[19]34, 6-26- [19]38, 9-8-[19]38, VI-6-[19]48, VI-30-[19]49 (2, MCZC; 8, OSAC0000551345–OSAC0000551348, OSAC0000551350[ Fig. 40a ]–OSAC0000551353); Pacific City, VII-18-1952, K. M. and D. M. Fender (1, OSAC0000551354); Sandlake, 6-13-1939, K. M. and D. M. Fender (3, OSAC0000551355, OSAC0000551356, OSAC0000551360); Seaside, Sept-15-1932 (1, OSAC0000551358); Tillamook, VIII-15-1937, Dan Bonnell (2, OSAC0000551349, OSAC0000551357); Woods, 7-25-1940, K. M. and D. M. Fender (1, OSAC0000551361). Washington: 27-VII-2-VIII-[20]02, P. M. Hammond/ Ocean shores, dunes, beach/ Hammond, P. M., BMNH(E) 2002-133 (1, BMNH); Gray’ s Harbor Co., Grayland, June-3-1951, Ruth Beresford (1, OSAC0000551337); Capalis [sic] Beach, VI- 13-[19]36/ Van Dyke collection (1, CASC); Twin Harb. Beach St. Pk., July-10-1949, M. H. Hatch (7, OSAC0000551328–OSAC0000551334); Pacific Co., Long Beach, July-4-1931, VII-15-[19]36 (1, OSAC0000551338; 6, CASC); Ilwaco, July 1918, O. E. Miner (1, WSUC); Ocean Park, June- 19-1930, July-4-1935 (5, OSAC0000551339, OSAC0000551340,OSAC0000551342–OSAC0000551344); Ocean Park, July-9-1948, R. P. Owen (1, OSAC0000551341); Ocean Park, July-12-1954, Helen B. James (1, WSUC); Ocean Park, June- 22-1956, June-23-1956 [ Figs. 40d, e ], June-29- 1957, M. T. James (23, WSUC); Seaview, July-2- 1929, July-18-[19]35 (2, OSAC0000551335, OSAC0000551336).

NEW WORLD SPECIES FORMERLY IN

HEMICREPIDIUS

Most of the Central American species of Hemicrepidius are moved herein to Athous . No Central American species has all of the diagnostic features of Hemicrepidius . No species has apico-ventral membranous lobes on the tarsi, although tarsomere 3 is expanded ventrally ( Figs. 4e–g ). No species has medial antescutellar notches ( Fig. 4h ). Most importantly, the internal female genitalia ( Figs. 8e–g ) are very different from those of Hemicrepidius ( Figs. 7a–e ), but match those of Athous vittatus , the type species of Athous ( Leseigneur 1972; personal examination). The ventrally expanded tarsomere 3 is similar to the condition seen in many Athous species , such as Athous subfuscus (Müller, 1764) . In his examination of the fauna of France, Leseigneur (1972: 193) stated that A. subfuscus had similar internal female genitalia to A. vittatus .

Preliminary molecular data ( Figs. 1–3 ) also support removal of these from Hemicrepidius . Hemicrepidius , as currently defined, is not known to occur in the Neotropics, including the West Indies.

Athous acuminatus ( Champion, 1896) (Hemicrepidius) , new combination

Athous bivittatus ( Champion, 1896) (Hemicrepidius) , new combination

Athous candezei ( Champion, 1896) (Hemicrepidius) , new combination

Athous chicamensis Etzler , new replacement name zoobank.org/ urn:lsid:zoobank.org:act:A15154C7-6288-4E80-B59A-5A703B82D9F7 Replacement name for Hemicrepidius biformis

Champion, 1896, a secondary homonym of Athous

biformis Redtenbacher, 1859, currently a synonym

of Athous zebei Bach, 1852 . Named for Chicam,

Guatemala, where this species is found.

Athous chiriqui Etzler , new replacement name zoobank.org/ urn:lsid:zoobank.org:act:47404311-235F-4E12-9EEF-97D951B5A506 Replacement name for Hemicrepidius long-

icornis Champion, 1896, a secondary homonym of

Athous longicornis Candèze, 1865 . The name is a

noun in apposition for the Chiriqui province in

Panama, where two of the type specimens were

collected.

Athous consanguineus ( Champion, 1896) (Hemicrepidius) , new combination

Athous consobrinus ( Champion, 1896) (Hemicrepidius) , new combination

Athous cruciatus ( Champion, 1896) (Hemicrepidius) , new combination

Athous deceptor ( Candèze, 1863) (Asaphes) , new combination, status resurrected

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

d) Dorsal view, e) Ventral view. Scale bars: 5 mm (a, b), 1 mm (c–e).

Asaphes longicollis Candèze, 1863 , reduced to synonymy with Athous deceptor , the next available name, to prevent secondary homonymy with Elater longicollis Olivier, 1790 , currently a synonym of Athous bicolor (Goeze, 1777) , and sometimes recognized as a valid species in Athous . Champion (1896) used this name over A. deceptor when he synonymized the two species.

Athous flavipes ( Candèze, 1863) (Asaphes) , new combination

Athous germanus ( Champion, 1896) (Hemicrepidius) , new combination

Athous instabilis ( Candèze, 1863) (Asaphes) , new combination

Asaphes leucostigma Candèze, 1863 , synonym of A. instabilis

Athous lecontei (Candèze, 1889) (Hemicrepidius)

Athous nitidus ( Champion, 1896) (Hemicrepidius) , new combination

Athous parvulus ( Champion, 1896) (Hemicrepidius) , new combination

Athous patruelis ( Champion, 1896) (Hemicrepidius) , new combination

Athous pictipes (Chevrolat, 1843) (Agrypnus) , new combination

Pomachilius longipennis ( Candèze, 1863) (Asaphes) , new combination

Sharon amoenus ( Philippi, 1861) (Asaphes) (moved in Arias and Elgueta 2015)

LOCALITY INFORMATION FOR NON-NORTH

AMERICAN HEMICREPIDIUS IN MOLECULAR

TREE

Athous deceptor , or species near Guatemala: Dept. Zacapa Sierra de los Minas; “El Naranjo” S slope below Sab Lorenzo Mine , 15.0733°, -89.6848°; 1600-1700m, 21-24-V-2010; oak forest, P. Skelley, flight intercept trap (1, FSCA; DNA— MTEC023726 ) .

Figures

Fig. 9. Morphological character measurements. Hemicrepidius bilobatus: a) Measurement of pronotal hind angle in dorsal view. H. tumescens: b) Measurement of pronotal hind angle in ventro-lateral view, d) Measurements of aedeagus. H. memnonius: c) Location of ventral setal patch (vsp) on paramere. L = length, w = width. Not to scale.

Fig. 40. Hemicrepidius tumescens. Dorsal habitus: a) Male, b) Female. c) Pronotum, lateral view. Male genitalia:

Fig. 41. Distribution maps of examined Hemicrepidius specimens (blue/gray dots). a) H. soccifer, b) H. tumescens.

Fig. 6. Morphological characters of Hemicrepidius. H. memnonius: a) Mesothorax, ventral view, b) Crenulate basal carina on elytra (arrow), c) Detail of subumbilicate punctures, lateral edge of pronotum, e) Microserrations on sides of abdominal ventrites, arrow. H. hemipodus: d) Simple basal carina on elytra (arrow).

Fig. 4. Morphological characters of Hemicrepidius and select genera. Dipropus: a) Head, anterior view; c) Pronotum, lateral view. H. memnonius: b) Head, anterior view; h) Pronotum, dorsal view. Athous instabilis: d) Pronotum, dorsal view; g) Tarsus, lateral view. A. vittatus: e) Tarsus, dorsal view; f) Tarsus, lateral view. Images not to scale. asn = antescutellar notch; fc = frontal carina; fcr = frontoclypeal region; hac = hind angle carina; il = impunctate line; lat = lateral area of pronotum; lpc = lateral pronotal carina; pd = pronotal disc; sln = sublateral notch.

Fig. 8. Hemicrepidius external female genitalia and Athous internal female genitalia. H. memnonius: a) Ovipositor, b) Ovipositor, detail of apex, c) Tergite VIII, d) Sternite VIII. A. instabilis: e) Dorsal, f) Lateral. Costa Rican Athous species: g) Lateral. bc: bursa copulatrix; cg: colleterial gland; co: entrance to common oviduct; te: tubular extension; v: vagina. Not to scale.

Fig. 7. Female internal genitalia. Hemicrepidius memnonius: a) Dorsal, b) Lateral, c) Ventral. H. soccifer: d) Dorsal, e) Lateral. bc: bursa copulatrix; cg: colleterial gland; co: entrance to common oviduct; te: tubular extension; v: vagina; spth: spermatheca (shrunken). Not to scale.

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.