Lissominae Laporte, 1835

Johnson, Paul J., 2021, Lissomus cacique, a Remarkable New Species of Click Beetle (Coleoptera: Elateridae) from Cloud Forests in Costa Rica and Panama, with Notes on Lissomus flavipennis Guérin-Méneville, The Coleopterists Bulletin 75 (3), pp. 599-611 : 600-603

Lissominae Laporte, 1835 Lissomini Laporte, 1835

Lissomus Dalman, 1824

Lissomus Dalman 1824: 13 .

Type species: Lissomus punctulatus Dalman 1824: 14 , designation of Fleutiaux 1947: 138.

Lissomus ; Eschscholtz 1829: 31, 1836: table; Latreille 1829a: 452, 1829b: 452, 1831: 342, 1832: 317, 1834a: 135, 1834b: 426, 1849: 510; Dejean 1833: 84, 1836: 95; Lacordaire 1830: 242, 1857: 135; Laporte de Castelnau 1835: 178, 1836: 260, 1840: 229; Chevrolat 1835: 194, 1849: 404, 1868: 147; Brullé 1837: 247; Germar 1839: 195; Voigt 1839: 89; Drapiez 1839: 432, 1853: 452; Guérin-Méneville 1838: 260, 1844b: 41; Blanchard 1846: 145; Bonvouloir 1859: 97, 1860: 366; Jacquelin du Val 1859: 110; LeConte 1859: 614; Schiödte 1866: 333; Gemminger and Harold 1869: 1459; Fleutiaux and Sallé 1889: 405; Heyne and Taschenberg 1905: 147; Fleutiaux 1911: 237; Schenkling 1928: 12; Blackwelder 1944: 304; Yensen 1975: 126; Costa et al. 1988: 154, 2010: 91; Aguirre-Tapiero 2009: 31; Aguirre-Tapiero et al. 2010: 20; Aguirre-Tapiero and Johnson 2014: 6; Johnson and Chaboo 2015: 271; Johnson et al. 2018: 82; Kundrata et al. 2019a: 124, 2019b: 5; Kirmse and Johnson 2020: 5.

Lissode Eschscholtz 1829: 31 (as synonym, not available, ICZN Art 11.6).

Lissodes Latreille 1829a: 452 (as synonym, not available, ICZN Art 11.6), 1829b: 452, 1831: 342, 1832: 317, 1834a: 135, 1834b: 426, 1849: 510; Laporte de Castelnau 1835: 178, 1840: 229; Voigt 1839: 89; Lacordaire 1857: 93; Bonvouloir 1859: 97; Gemminger and Harold 1869: 1459.

Cymbium Gerstäcker 1860: 136 (not Cymbium Röding 1798: 151 ; Gastropoda ) (as synonym, not available, ICZN Art. 11.6).

Pseudelater Fleutiaux 1899: 161 (as synonym, not available, ICZN 1999:Art. 11.6) (not Pseudelater Miwa 1934: 20 ; type species Elater habunensis Miwa 1934: 20 , original designation).

Lissernus Fleutiaux 1911: 263 (misspelling).

Diagnosis [Adapted after Bonvouloir (1859) and Gerstäcker (1860)]. Body moderate in length, about 6–16 mm; dorsum shallowly to strongly convex, venter shallowly convex. Integument smooth, shining, finely to moderately punctured; each puncture with either a minute fine seta and surface appearing glabrous, or seta long, thin, hair-like. Head depressed to shallowly impressed on frons, frontal margin a thick medial intra-antennal pad and a strong subantennal carina each side. Mandibles stout, acutely bidentate apically. Antennal insertion fossa deep, transverse; antenna with antennomere 1 robust, subcylindrical to subangular, coarsely punc- tate, slightly arched; antennomere 2 subcylindrical to subserrate; antennomeres 3–10 moderately to strongly serrate, antennomere 11 ovoid to lachrymiform. Pronotum trapezoidal, shallowly to moderately convex, sparsely to moderately densely punctured; anterior angles often slightly reflexed; hind angles with dorsal carina extended anterad and forming a longitudinal bulge before anterior angle. Prosternum longitudinally convex anterad of procoxal cavities, shallowly to moderately transversely arched at midlength, with anterior lobe not to shallowly deflexed; intercoxal process with apex broadly to narrowly rounded, postcoxal portion slightly expanded laterally, and with strong dorsal extension; lateral margins simple to shallowly sulcate. Hypomeron deeply excavate posteriorly to receive front legs, posterior margin a thin perpendicular lamelliform wall; pro- notosternal margin deeply excavate to receive antenna. Scutellar shield transverse, ovoid to subpentagonal with obtuse angles; dorsum flat to shallowly convex, sparsely punctured, shining. Elytra moderately to strongly convex, occasionally depressed discally, often gibbous; striae 1–8 of fine to moderately-sized serial punctures; intervals flat to weakly convex; stria 9 coarsely punctured to sulcate; apices conjointly rounded, often deplanate. Mesoventrite with deep median fossa, broadly U- to V-shaped, walls planar with metaventrite to below or slightly elevated; anterior angles formed as crural impressions to receive mesothoracic legs, usually with an arcuate post-coxal carina from mesocoxal margin directed laterally, reaching partly or com- pletely to lateral margin. Tarsomeres 1–4 with pro- gressively enlarging ventral membranous lobes, rarely with tarsomere one with densely setose ventral pad. Pretarsal claw asetose. Abdomen with 5 ventrites, ventrite 1 with elytral locking flange laterally, ventrite 5 broadly rounded apically. Male with sternite 8 rounded apically, V-shaped basally; sternite 9 membranous discally, truncate to shallowly emarginate apically; tergite 9 with deep V-shaped emargination; tergite 10 subtruncate apically; aedeagus trilobed with median lobe and parameres artic- ulating against basal piece; median lobe variably attenuate; parameres sagittate, hooked or truncate at apex. Female with sternite 8 narrowly rounded apically, broadly U-shaped basally, with long, narrow strut; ovipositor with long, flattened, darkly sclerotized paraprocts; gonocoxites narrowly subtriangu- lar, usually with short subapical dorsolateral stylus bearing a long apical seta. Internal genital tract with bursa bulbous, with corrugated surface, 4 darkly sclerotized internal armatures, tubular colleterial glands from vaginal-bursal juncture, and spermathe- ca lateral, thin, corrugated, twisted.

Notes. The names Lissode ( Latreille 1825: 248), and Lissome and Lissomes ( Brullé 1837: 427; Latreille 1829a: xxiii, 452, 1829b: xxiii, 452, 1834a: 135), are French vernacular and are not available genus names. Eschscholtz (1829) appears to have treated “ Lissode ” as a genus and synonym of Lissomus in the same manner as other names for other taxa; this treatment is accepted here. As such, Lissode Eschscholtz is not an available name ( ICZN 1999: Art. 11.6). Earlier, Lissodes Berthold (1827: 335) was presented as a genus name, but there were no assigned or indicated species, hence this is not an available name ( ICZN 1999:Art. 12.2.5) for purposes involving Lissomus nomenclature. Treatment of Lissodes as a synonym of Drapetes Megerle by Smetana (2007) is a nomenclatural issue best treated elsewhere.

LISSOMUS CACIQUE Johnson , new species zoobank.org/ urn:lsid:zoobank.org:act:23443A9F-DA2D-47F0-B08B-41128E2268B9 ( Figs. 1, 3, 5, 7–9 )

Diagnosis. Lissomus cacique is immediately rec- ognized by the combination of larger size, subparallel dorsal silhouette at midlength, strongly convex dorsum ( Fig. 3 ), shallowly subsulcate elytral striae, and orange-red elytra contrasting with black body. This species is superficially similar to L. flavipennis from Mexico, and will key to this species in Bonvouloir (1859) and Gerstäcker (1860). Both species share the traits of an arcuate post-coxal carina ( Fig. 8 ) delimiting the posterior margin of the crural impression for the mesothoracic leg; the anterior pronotal angles appearing constricted at a smooth bulging termination of a carina; and the elytra predominately of a translucent yellow-brown to orange-red color (post-mortem vs. alive) with narrow black margin at apices, strikingly different from the remainder of the body, with visible trabeculae, and with a deep stria 9 sulcus usually separated from humeral sulcus by a trans-sulcal bridge; elytral margin carinate only at base, generally rounded, smooth, polished, contiguous with epipleuron. In contrast, L. flavipennis is proportionately shorter, more oval in outline, and the elytral striae and intervals form a continuous smooth surface.

Description. Body length 13.9–14.8 mm, width 6.0– 6.1 mm across humeri, 6.6–6.9 mm across elytral midlength, oblong, subparallel to el- liptical in dorsal silhouette ( Fig. 1 ), dorsum strongly convex ( Fig. 3 ). Integument black thoughout, except elytra rufotestaceous to testaceous, with black margins and suture near apices ( Figs. 1, 3 ). Pubescence fine, pale flavous, moderately long, minute on elytra. Head coarsely punctured, punctures large, deep, umbilicate, approximate; frons shallowly impressed. Antennomeres 2–3 subserrate, subequal in length, 4–10 strongly serrate, 11 lachrymiform, 2–11 length ratio 1.0:1.0:1.7:1.6:1.6:1.6:1.7:1.7:1.7:2.6. Pronotum broadly trapezoidal, coarsely punctured as on head, punctures approximate discally to sub- rugose laterally; median line polished, with irregular margins; posterolateral area shallowly impressed. Anterior margin bisinuate; anterior angles rounded, slightly reflexed. Lateral margin sinuate, joining medial hypomeral margin before anterior angle; pronotum appearing constricted at anterior quarter. Hind angle dorsal carina long, reaching two-thirds the distance from hind angle apex to anterior margin, close to lateral margin, terminating on lateral dilation of pronotum. Scutellar shield shallowly convex, nearly flat, sparsely punctured. Elytra appearing glabrous, setae fine, minute, pale. Dorsal color translucent dull orange-red, revealing narrow and transverse linearly arranged trabeculae beneath striae. Humerus a shallowly to moderately domed boss, extended to fit over pronotal margin, subtend- ed by deep groove. Striae 1–8 finely, shallowly impressed to subsulcate; stria 9 deeply sulcate basally from near humerus, separated from subhu- meral groove by a smooth bridge, sulcus becoming shallower apically. Intervals shallowly convex, finely, sparsely, serially punctured. Costal margin smooth, rounded and confluent with epipleuron; lateral carina short, at base only. Prosternum with intercoxal process attenuate posteriorly, evenly convex, midline smooth and polished. Mesoventrite with arcuate carina delimiting crural impression for mesothoracic leg ( Fig. 8 ). Ventral sclerites moderately to coarsely punctured, slightly denser on ab- dominal ventrite 5. Metatarsomere length ratio 1.0:0.3:0.3:0.3:1.1. Aedeagus ( Fig. 5 ) with basal piece 0.42–0.43 of total length; median lobe narrow, evenly attenuate, apex acute, 0.72–0.74 of total length; parameres gradually narrowing apically, 0.76 of total length; apices obtuse, with acute subapical lateral spine, 0.06 of paramere length. Females not distinguishable from males without examination of terminal sclerites or genitalia. Gonocoxites and internal genital tract ( Fig. 7 ) as per genus diagnosis.

Type Material. Holotype, female: “Buen Amigo, San Luis, Monteverde, Pr. Punta., COSTA RICA, 1000–1350m, 20 MAR-14ABR 1995, M. Segura, L N 250850 449250 #4410/INBIOCR002191821”. This specimen will be returned to the INBio collection, presently managed by the Museum Nacional de Costa Rica , San José, Costa Rica.

Paratypes (28; 5 females, 5 males available for dissection). COSTA RICA: Cartago Province: Parque Nacional Tapanti, Orosi, 1,500 m, 23-AUG-1991, R. Coto (1, INBIO / MNCR) . Guanacaste Province: Estac. Cacao , 1000–1400m, SW side Volcan Cacao, Guanac. Pr., Mar 1988, GNP Biod. Sur., 32300, 375700/INBIO CRI00050623 (1, INBIO / MNCR); same, Mar 1988, GNP Biod. Sur., 32300, 375700/CRI000050620 (1, INBIO / MNCR); same/CRI000050626 (1, INBIO / MNCR); same, Apr 1988 /INBIO CR000035273 (1, INBIO / MNCR), same/CRI000127682 (1, INBIO / MNCR); same, M. Espinoza /CRI000036398 (1, INBIO / MNCR); same/ CRI000036130 (1, INBIO / MNCR); Malaise Tp, 1988–1989, GNP Biodiv. Survey 323300, 375700/ CRI000071599 (1, INBIO / MNCR); A.C.G. Liberia, Pque Nal Gte. Est Cacao . 1000–1150m, 9 MAR 1988, Espinoza Manual, L _N_323150_375500 #52583/INB0003354495 (1, INBIO / MNCR); Estac. Menog , 1100m, SW side Volcan Cacao, Guanac. Pr., Feb 1989, GNP Biodiversity Survey, W85 28′10′′, N10 55′43′′ /CRI001038133 (1, INBIO / MNCR) . Puntarenas Province: Buen Amigo, San Luis Monteverde, A.C. Arenal, Prov. Punta , 1000–1350 m, May 1994, Grace Fuentes, L N 250850_449250 #2927/CRI001870832 (1 INBIO / MNCR); same, ABR 1995, Z. Fuentes, L N 250820 449250 #4801/ CRI002202986 (1, INBIO / MNCR); Cerro Amapola, San Luis, Monteverde , Prov. Punta , 1000–1350m, 23 Feb 1994, Z. Fuentes, L N 250850_449250 #2620/CRI001981686 (1, INBIO / MNCR); Monte Verde, Pens [ion] Quetzal , v-18-1988 , 1380 m, B. Ratcliffe & M. Jameson (1, UNLC) ; Monteverde de Puntarenas 1959, has. Palmer /M. Robinson collection 1959 (1, FSCA) , 1-3.vi.1978, E. Giesbert Coll. (1, FSCA) , 11-14.iv.1981, E. Giesbert Coll. (1, FSCA) , 23-27.v.1987, E. Giesbert Coll. (1, FSCA) ; San Luis, 1040 m, R. B. Monteverde, Prov. Punt. , Abr 1993, Z. Fuentes, L-N250820,449250/INBIO CR001325792 (1, MUCR) ; Monteverde, v.26– vi.3.84, E. Riley, D. Rider, D. Ledoux (2, TAMU) ; C. R. Punt. ACLA. R.P. Z.P.T., Send. al Salto a 4.5 km, NE de Progreso , 1600m, 18-enero-1998, A. Picado Col.en Tronco podsido, LS-320150/599000E/ CRI002607764 (1, INBIO / MNCR); C. R. Prov. Punt. Est. Pittier Send. a Río Canasta, 1.4 Km NNO. de la Estación 1750-1800m, 26–27 MAR 1998, M. M. Morago, Red Mariposera, LS _332100_576800 #50835/INB0003014982 (1, INBIO / MNCR); Est. Biol. Las Alturas , 1500m, Coto Brus, Prov. Punt, M. Zumbado, Ene 1992, L-S-322500, 591300/ CRI000339475 (1, INBIO / MNCR) . PANAMA: [Chiriquí Province:] Hartmann’s Finca , 19.v.1996, R. Turnbow (1, FSCA) .

Additional Specimen as Image. COSTA RICA, [Puntarenas Province] Monteverde , April 2013 ,

Figures

Figs. 1–7. Lissomus species. Adult male, dorsal aspect: 1) L. cacique, new species; 2) L. flavipennis. Adult male, lateral aspect: 3) L. cacique, new species (photo by H. Lezama, used with permission); 4) L. flavipennis. Aedeagus, dorsal aspect: 5) L. cacique, new species; 6) L. flavipennis. 7) L. cacique, new species, female internal genital tract.

Fig. 8. Lissomus cacique, new species, ventral oblique aspect, arrow pointing to the arcuate postcoxal carina.

Fig. 9. Satellite image of Costa Rica and Panama with five Lissomus cacique, new species sites pinned. Two sites have multiple nearby collections as indicated.