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Isarthrus striatus Otto, Muona and Córdoba-Alfaro, new species Figures 35–36 Diagnosis. The new eucnemid species can be readily distinguished from any macraulacine members of Eucnemidae present in Costa Rica by the presence of an interantennal carina, a wide lateral antennal groove which is widest in the middle, simple tarsal claws and dark colored dorsum with reddish-brown antennae and legs. The new species differs from the Nearctic Isarthrus calceatus LeConte by its antennal grooves widest in the middle, striate elytra and shorter flagellomeres II–III. Isarthrus calceatus have parallel-sided antennal grooves, poorly developed striae on the elytra, and slightly longer flagellomeres II–III. Type material. Female holotype: “ COSTA RICA: Prov. Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; 24 Feb 2004; INBio-OET-ALAS transect” (“24 Feb” handwritten on label) / “ 24 Febrero 2004; Malaise; M/26/768” / “ HOLOTYPE:; Isarthrus; striatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Paratypes. 4 ♀♀: COSTA RICA: 1 ♀, “COSTA RICA: Heredia; Pr: La Selva Bio. Sta.; 3 km s. Pto. Viejo; 10°26′N 84°01′W ” / “ ii–iv.1993, P. Hanson, Malaise Trap ” / “ PARATYPE:; Isarthrus; striatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♀, “COSTA RICA: Heredia; Pr: La Selva Bio. Sta.; 3 km s. Pto. Viejo; 10°26′N 84°01′W ” / “ iv–v.1993, P. Hanson, Malaise Trap ” / “ PARATYPE:; Isarthrus; striatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♀, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/04/111; 01 Junio 1993; Bosque primario” / “ PARATYPE:; Isarthrus; striatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♀, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/10/172; 03 Augosto 1993; Bosque primario” / “Collection of the Global; Eucnemid Research Project; (Robert L. Otto)” (green framed white label) / “ PARATYPE:; Isarthrus; striatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (GERP). Paratypes are to be deposited in INBC and GERP. Description. Female holotype: Length, 8.5 mm. Width, 2.5 mm. Body elongate; dark brownish-black; antennae and legs dark reddish-brown; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 35). Head: Surface somewhat shiny, subspherical with crowded punctures; frons convex with slight circular impression above interantennal carina, poorly developed median carina present; interantennal carina present; apical margin of frontoclypeal region weakly trilobed, more than 2 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Filiform from flagellomeres I–IX, attaining about 1/3 the length of the body; flagellomere I longer than wide, longer than II; flagellomeres II–III each slightly longer than wide, sub-equal, each shorter than I and IV; flagellomeres IV–V each longer than wide, sub-equal, longer than III, shorter than VI; flagellomeres VI–VIII each longer than wide, sub-equal; flagellomere IX slightly longer than VIII. Pronotum: Surface somewhat shiny, with deep, crowded punctures; quadrate, with well-developed, sharp hind angles; sides parallel-sided at basal 1/2, arcuate at apical 1/2; disc convex, slightly depressed near base above scutellar shield; base sinuous. Scutellar shield: Quadrate, sub-triangular, shallowly punctate, setose and distally rounded. Elytra: Very shallow striae present; interstices slightly elevated; surfaces shiny, closely punctate. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV excavated; metatarsomere V elongate with simple claws. Venter (Figure 36): Closely punctate, with short, recumbent silver-colored setae; hypomeron with basally opened, lateral antennal grooves, widest in the middle; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Variations. Four female paratypes were examined. These paratypes measured 7.0– 8.5 mm long and 2.0– 2.5 mm wide. One female paratype is as large as and just as wide as the holotype. The remaining three paratypes are shorter and narrower than the holotype. Median carina on the frons are evident in two paratypes and absent in the other two paratypes. Distribution. This eucnemid species is known from five specimens taken from La Selva within the province of Heredia. Biology. Two paratypes were taken from a primary forest. Three specimens were taken from a Malaise trap. Larvae and pupae are unknown. Etymology. The specific epithet is derived from the presence of well-developed striae on the elytra which serves as one of the best feature to distinguish the new species from the Nearctic I. calceatus., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 18-20

Deltometopus bicolor Otto, Muona and Córdoba-Alfaro, new species Figures 64–66 Diagnosis. The new eucnemid species is very similar to D. fastidiosus Bonvouloir. Deltometopus bicolor can be distinguished from D. fastidiosus by the presence of interrupted interantennal carina at the base of the frontoclypeal region. Dromaeolus fastidiosus have a complete interantennal carina.The new species is further distinguished from D. foveolatus by its elongate, less cuneiform habitus and absence of sulcus along lateral margin of the pronotum. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/13/028; 03 Marzo 1993; Bosque secundario” / “ HOLOTYPE:; Deltometopus; bicolor ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Female allotype: “ COSTA RICA, Heredia; Est.Bio. La Selva, 50–; 150 m, 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/13/028; 03 Marzo 1993; Bosque secundario” / “ ALLOTYPE:; Deltometopus; bicolor ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Holotype and allotype are to be deposited in INBC. Paratypes. 2 ♂♂, 1 ♀: COSTA RICA: 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/09/072; 15 Abril 1993; Bosque secundario” / “ PARATYPE:; Deltometopus; bicolor ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/11/341; 15 Febrero 1994; Bosque secundario” / “Collection of the Global; Eucnemid Research Project; (Robert L. Otto)” (green framed white label) / “ PARATYPE:; Deltometopus; bicolor ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (GERP); 1 ♀, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/02/400; 17 Julio 1995; Bosque secundario” / “ PARATYPE:; Deltometopus; bicolor ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC). Paratypes are to be deposited in INBC and GERP. Description. Male holotype: Length, 2.5 mm. Width, 1.0 mm. Body elongate; dark black; antennae brown; femur and tibiae brown, tarsi brown; head, pronotum and elytra clothed with short, recumbent whitish setae (Figure 64). Head: Surface densely punctate, somewhat shiny, subspherical; frons convex, without fovea or ridge; apical margin of frontoclypeal region rounded, more than 2 times wider than base; interantennal carina incomplete; mandibles stout, bidentate, densely punctate. Antenna: Serriform from flagellomeres I–VIII, attaining about 1/3 the length of the body, just beyond pronotal hind angles; flagellomere I longer than II; flagellomeres II–III sub-equal, slightly longer than wide; flagellomeres IV–VIII sub-equal, each slightly larger than III, slightly longer than wide; flagellomere IX simple, slightly longer than VIII. Pronotum: Surface somewhat shiny, densely punctate; quadrate, with moderate, sharp hind angles; lateral sides sinuous, apically narrowed, arcuate; disc convex without fovea or ridge; base sinuous. Scutellar shield: Transverse, sub-triangular, shallowly punctate and distally rounded. Elytra: Striae delicately indicated on humeri, indistinct elsewhere; interstices flattened; surfaces shiny, transversely rugose at basal 1/4, dense, crowded, shallow punctures present elsewhere. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV short, excavate-emarginate; metatarsomere V elongate with simple claws. Venter (Figure 65): Closely punctate, with short, recumbent white-colored setae; hypomeron with basally widened, basally opened, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Female allotype (Figure 66): Length, 4.0 mm. Width, 1.0 mm. Flagellomeres serriform, about 1/3the length of the body; flagellomere I longer than wide, longer than II; flagellomere II–IV sub-equal, quadrate; flagellomeres V–VIII each slightly longer than IV, quadrate, sub-equal. Frons impressed above frontoclypeal region. Variations. Three paratypes were examined. Two male paratypes measured 3.0 mm long and 1.0 mm wide. One female paratype measured 4.0 mm long and 1.0 mm wide. All paratypes are longer than and just as wide as the holotype. Both male paratypes have a more pronounced impression on the frons just above the frontoclypeal region. Frons on the female paratype are simple, without any indications of impression above the frontoclypeal region. Pedicel and flagellomere I are yellowish in one male paratype. The remaining two paratypes have antennal colorations similar to the holotype. Habitus in all paratypes are just as dark as the holotype. Striae at the humeri are more pronounced in one of the male and female paratypes compared to the holotype. Striae in the other male paratype are similar to the holotype. Distribution. This eucnemid species is known from five specimens taken from a single location in four different collection events at La Selva within the province of Heredia. Biology. All specimens in the type series were taken from a second growth forest. Larvae and pupae are unknown. Etymology. The specific epithet is derived from its black habitus along with dark brown legs and antennae. Tribe Nematodini Leiler, 1976 Diagnosis. Mandibles short, with ventral secondary tooth, without expanded lateral surfaces; prothoracic tibiae with one apical spur; male prothoracic tarsomere I with basal sex combs; tarsomere IV usually bilobed; lateral sides of mesothoracic and metathoracic tibiae variable, either with setae and simple spines or with setae and transverse rows of spine combs; hypomeron with or without antennal grooves; prothoracic sternal peg high, either truncated or excavated; median lobe without dorsal basal struts, fused with lateral lobes, distinct, with narrowly and deeply bifurcate apex; bursa divided, simple; spermatheca sclerotized, divided and U-shaped (Muona 1993; Otto 2017c)., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 30-31, {"references":["Muona J. 1993. Review of the phylogeny, classification and biology of the family Eucnemidae (Coleoptera). Entomologica Scandinavica Supplement 44: 1 - 133.","Otto RL. 2017 c. Descriptions of six new species of false click beetles (Coleoptera: Eucnemidae: Macraulacinae) with new identification keys for one tribe and two genera. Insecta Mundi 0558: 1 - 19."]}

Onichodon rufus Otto, Muona and Córdoba-Alfaro, new species Figures 33–34 Diagnosis. The smaller size along with the reddish coloration with its habitus will distinguish the new species from O. confluentus new species and any other misplaced members of Fornax present in Costa Rica. Type material. Female holotype: “ COSTA RICA: Heredia; pr: La Selva Bio. Sta.; 3 km S Pto. Viejo; 10°26′N 84°01′W ” / “ 25.IV.1990; H.A. Hespenheide ” (“25”, “IV” and “0” handwritten on label) / “ HOLOTYPE:; Onichodon; rufus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Description. Female holotype: Length, 6.0 mm. Width, 1.5 mm. Body elongate; dark reddish; antennae and legs dark reddish; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 33). Head: Surface somewhat shiny, subspherical with crowded punctures; frons convex without carina or fovea; interantennal carina present; apical margin of frontoclypeal region weakly trilobed, more than 2 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Weakly serriform from flagellomeres I–IX, attaining about 1/3the length of the body, just beyond pronotal hind angle; flagellomere I longer than wide, longer than II; flagellomeres II–VIII each slightly longer than wide, sub-equal; flagellomere IX slightly longer than VIII. Pronotum: Surface somewhat shiny, with deep, crowded punctures; quadrate, with well-developed, sharp hind angles; sides parallel-sided at basal 2/3, arcuate at apical 1/3; disc convex, without carina or fovea; base sinuous. Scutellar shield: Slightly longer than wide, sub-triangular, shallowly punctate, setose and distally rounded. Elytra: Very shallow striae present; interstices slightly elevated; surfaces shiny, transversely rugose through much of the elytra. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV excavated; metatarsomere V short with basally toothed claws. Venter (Figure 34): Closely punctate, with short, recumbent yellowish colored setae; hypomeron with basally opened, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Distribution. This eucnemid species is known from the holotype taken from La Selva within the province of Heredia. Biology. No biological information is known for the new species. Larvae and pupae are unknown. Etymology. The specific epithet is derived from its overall reddish coloration of its habitus., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 17-18

Dromaeolus holdridgei Otto, Muona and Córdoba-Alfaro, new species Figures 61–62 Diagnosis. The new eucnemid species can be readily distinguished from any ornate members of Dromaeolus present in Costa Rica by the presence of interantennal carina, very short, median carina on the frontoclypeal region and short, median carina present on the frons except D. herediensis new species. Narrower silver-colored band on the elytra along with the absence of silver-colored setae at elytral apices will distinguish the new species from D. herediensis new species. Type material. Female holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/01/136; 01 Julio 1993; Parcelas sucesionales” / “ HOLOTYPE:; Dromaeolus; holdridgei ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Description. Female holotype: Length, 6.0 mm. Width, 1.5 mm. Body elongate; black; antennae black; femur and tibiae black, tarsi dark brownish-black; head and pronotum clothed with short, recumbent silver-colored setae more apparent at base and along lateral sides of pronotum, elytra with silver-colored setae more apparent along lateral sides at basal half, narrow band across the middle of each elytron and narrow band along elytral suture terminating at the middle band (Figure 61). Head: Surface dullish, subspherical with closely spaced punctures; frons convex with short median carina; frontoclypeal region with short, delicate median carina; interantennal carina present, sinuous; apical margin of frontoclypeal region weakly trilobed, more than 2 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Filiform from flagellomeres I–IX, attaining nearly 1/3the length of the body, never passed beyond pronotal hind angles; flagellomere I longer than wide, longer than II; flagellomere II quadrate, shorter than III; flagellomere III slightly longer than wide, shorter than IV; IV–VIII each sub-equal, longer than wide; flagellomere IX slightly longer than VIII. Pronotum: Surface shiny, with evenly dispersed punctures; slightly longer than wide, with well-developed, sharp hind angles; sides subparallel-sided, slightly arcuate near craniad; disc convex without shallow median groove or circular fovea; base sinuous. Scutellar shield: Elongate, sub-triangular, shallowly punctate, setose and distally rounded. Elytra: Very shallow striae indicated; interstices slightly elevated; surfaces shiny with transversely rugose to deep punctures at humeri, shallower punctures elsewhere. Legs: First tarsomere longer than the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV excavated; metatarsomere V short with simple claws. Venter (Figure 62): Closely punctate, with short, recumbent silver-colored setae; hypomeron with basally opened, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Distribution. This eucnemid species is known from a single specimen taken from La Selva within the province of Heredia. Biology. The holotype was taken from a successional plot. Larvae and pupae are unknown. Etymology. The specific epithet is dedicated in honor of Dr. Leslie Holdridge, an American botanist and climatologist who was the founding member of the La Selva Biological Station established in 1968., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 28-29

Genus Quirsfeldia Cobos, 1964 Diversity and distribution. The small group consists of several species with two being present in the Amazonian region of South America, particularly in Brazil and a third species recorded for the first time in Central America. The new Central American species is only known from a single province in Costa Rica. Diagnosis. Apical margin of frontoclypeal region fairly evenly rounded and less than twice as wide as the distance between antennal sockets; poorly indicated notosternal antennal grooves present, caudally obliterated; elytral apices meeting tightly together; metacoxal plates parallel-sided, medially less than 2.5 times wider than laterally; frons simple; last visible ventrite acutely produced; simple tarsal claws; lateral surfaces of meso- and metatibiae with setae only (Muona 2011). The genus is very similar to Adelothyreus Bonvouloir and can be distinguished by the apical margin of the frontoclypeal region being less than twice as wide as the base. Adelothyreus has the apical margin of the frontoclypeal region more than twice as wide as the base., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on page 10, {"references":["Muona J. 2011. Eucnemidae. info Homepage. Available at http: // dol. luomus. fi: 8080 / cgibin / dol / dol _ homepage. pl (Last accessed 15 August 2019.)"]}

Lacus pectinatus Otto, Muona and Córdoba-Alfaro, new species Figures 23–26 Diagnosis. The new eucnemid species is distinguished from L. laticornis by the presence of broken setal patterns on the pronotum. Lacus laticornis have a more intact bands of setal patterns present on the pronotum. Punctate striae are present in the new species, whereas striae in L. laticornis is solid. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/02/065; 15 Abril 1993; Bosque secundario” / “ HOLOTYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Female allotype: “ COSTA RICA, Heredia; Est.Bio. La Selva, 50–; 150 m, 10°26′N 84°01′W; Apr 2000, INBio-OET” / “ 17 Abril 2000; Bosque secundario; M/19/746” / “ ALLOTYPE:; Lacus; pectinatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Holotype and allotype are to be deposited in INBC. Paratypes. 6 ♂♂, 3 ♀♀: COSTA RICA: Heredia Province: 1 ♂, “COSTA RICA: Heredia; Pr: La Selva Biol. Sta.; 3 km S Pto Viejo; 10°26′N 84°01′W ” / “ 12 FEB 1993; Bosque secundario; M/00/011” / “ PARATYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/02/017; 02 Marzo 1993; Bosque secundario” / “ PARATYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♀, “COSTA RICA: Heredia; Pr: La Selva Biol. Sta.; 3 km S Pto Viejo; 10°26′N 84°01′W ” / “ 2 Mayo 1993; Bosque secundario; M/02/081” / “ PARATYPE:; Lacus; pectinatus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (INBC); 1 ♂ 1 ♀, “COSTA RICA: Prov. Heredia; Est. Biol. La Selva, 50–; 150, 10°26′N 84°01′W; 09 Marzo 2004, INBio-OET-ALAS transect” (“09 Marzo” handwritten on label) / “ 09 Marzo 2004; Malaise; M/25/777” / “ PARATYPE:; Lacus; pectinatus; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (either ♂ or ♀ printed behind species name on label) [yellow printed label] (INBC); 1 ♂, “COSTA RICA: Prov. Heredia; Est. Biol. La Selva, 50–; 150, 10°26′N 84°01′W; 21 Marzo 2004, INBio-OET-ALAS transect” (“21 Marzo” handwritten on label) / “ 21 Marzo 2004; Malaise; M/23/785” / “Collection of the Global; Eucnemid Research Project; (Robert L. Otto)” (green framed white label) / “ PARATYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label) (GERP); Puntarenas Province : 1 ♂, “COSTA RICA. Prov. Puntarenas; Golfito Poue Nal Corcovado Est.; Agujas, Cerro Rincón. 745 m 13 FEB; 2000. A. Azofeifa. Intersección; L S 275500 522000 #56525” / “ PARATYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro, Det. J. Muona; 2022 (yellow printed label) (INBC); PANAMA: Col ó n Province: 1 ♂, “IBISCA Project Panama; Colon, San Lorenzo NP; 27.3–7.3 2004” (‘27.3–7.3’ handwritten on label; black framed white label) / “IBISCA 2003-05; Morphospec. sort; Jürgen Schmidl; COEUCN 50” (‘50’ handwritten on label; black framed white label) / “ Photo PACOEUCN; #146” (‘146’ handwritten on label; black framed pink label) / “ PARATYPE:; Lacus; pectinatus ♂; Otto, Muona & Córdoba-Alfaro; Det. J. Muona; 2022” (yellow printed label) (JMC); Panamá Province: 1 ♀, “ Panama; Canal Zone” / “ PARATYPE:; Lacus, pectinatus ♀; Otto, Muona & Córdoba-Alfaro; Det. J. Muona; 2022” (yellow printed label) (JMC). Paratypes are to be deposited in INBC, GERP and JMC. Description. Male holotype: Length, 6.5 mm. Width, 2.0 mm. Body obtuse; uniformly dark brown with infuscate reddish basal margin of elytral humeri, venter reddish-brown; antennae and legs reddish-brown; head, pronotum and elytra clothed with short, recumbent yellowish setae; broken setal patterns present on the pronotum (Figure 23). Head: Surface densely punctate, somewhat dullish, subspherical; frons convex, without median carina or fovea above frontoclypeal region; apical margin of frontoclypeal region rounded, about 2 times wider than base; interantennal carina absent; mandibles stout, bidentate, densely punctate. Antenna: Pectinate from flagellomeres I–VIII, attaining about 1/3the length of the body, never beyond pronotal hind angles; flagellomere I slightly longer than II; flagellomeres II–VIII each sub-equal, slightly longer than wide; rami moderately elongate, arising near apices of flagellomeres I–VIII; flagellomere IX simple, slightly longer than VIII. Pronotum: Surface somewhat dullish, densely punctate; slightly transverse, with moderate, sharp hind angles; lateral sides strongly arcuate and narrowing towards craniad; disc convex with pair of shallow circular fovea; base sinuous. Scutellar shield: Elongate, sub-triangular, setose, shallowly punctate and distally rounded. Elytra: Distinctly, shallowly punctate striate present; interstices slightly elevated; surfaces shiny, transversely rugose at humeri, dense, crowded punctures present elsewhere. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV very short, excavated, slightly emarginated; metatarsomere V elongate with basally swollen, simple claws. Venter (Figure 24): Closely punctate, with elongate, recumbent yellowish setae; hypomeron with basally closed, wide, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Aedeagus (paratype) (Figure 25): Elongate, dorsally flattened, sinuate; median lobe free, short, apically rounded; parameres apically elongate, rounded, longer than median lobe, without lateral tooth; secondary lateral lobe absent; basal piece elongate, apically truncated. Female allotype (Figure 26): Length, 8.0 mm. Width, 2.5 mm. Antennae filiform, about 1/3the length of the body; flagellomere I longer than wide, slightly longer than II; flagellomere II longer than wide, much longer than III; flagellomeres III–VIII quadrate, sub-equal; left flagellomere IX missing; right antenna missing. Lateral side of elytra above epipleura infuscate reddish. Variations. Six paratypes were examined. Several additional included paratypes from the Osa Peninsula and Panama were not examined for variations. Four male paratypes measured 6.0–7.0 mm long and 2.0 mm wide. Two female paratypes measured 5.0–7.0 mm long and 1.5–2.5 mm wide. Two of the male paratypes have the same length and width as the holotype. One male is shorter than and just as wide as the holotype. One male is longer than and just as wide as the holotype. One female is shorter and narrower than the holotype. One female is larger and wider than the holotype. All four males have a more pronounced reddish coloration along the humeri region than the remaining areas of the elytra. The reddish area of the humeri in both females are more infuscate and blended with the surrounding dark brown coloration of the elytral region. All paratypes are slightly darker than the holotype. Distribution. This eucnemid species is known from eight specimens taken from La Selva within the province of Heredia, a single specimen each from the Puntarenas province and two sites in Panama. Biology. Three specimens were taken from a Malaise trap. Five specimens were taken from a secondary growth forest. Larvae and pupae are unknown. Etymology. The specific epithet is derived from the presence of pectinate antennae found in male specimens., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 12-14

Absensiugum brunneum Otto, Muona and Córdoba-Alfaro, new species Figures 37–40 Diagnosis. The new species differ from A. teres by its antennal segments; stouter, quadrate in the new species and longer than wide in A. teres. Additionally, habitus coloration will further distinguish the new species from A. teres; dark brown in the new species, reddish-brown in A. teres. Type material. Male holotype: “ COSTA RICA, Heredia; Est.Bio. La Selva, 50–; 150 m, 10°26′N 84°01′W; Apr 2000, INBio-OET” / “M/01/236; 15 Octubre 1993; Parcelas sucesionales” / “ HOLOTYPE:; Absensiugum; brunneum ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Female allotype: “ COSTA RICA, Heredia; Est.Bio. La Selva, 50–; 150 m, 10°26′N 84°01′W; Apr 2000, INBio-OET” / “M/12/390; 04 Abril 1994; Bosque primario” / “ ALLOTYPE: Absensiugum; brunneum ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Holotype and allotype are to be deposited in INBC. Description. Male holotype: Length 3.0 mm, width 0.75 mm. Body color uniformly dark brown (Figure 37). Head: Subspherical, without carina or fovea on frons; interantennal carina absent; surface dullish; punctures deep, closely spaced; eyes protuberant. Antennae: Dark reddish-brown; filiform, about 1/3 of body length; flagellomeres II–IV quadrate, sub-equal; flagellomeres V–VIII each slightly larger than IV, quadrate; flagellomere IX larger than VIII; lateral carina absent on flagellomeres II–VIII. Pronotum: Surface dullish, with somewhat elongate, yellow recumbent setae; punctures deep, closely spaced, almost rugose; longer than wide, with moderate, sharp hind angles; sides subparallel, sinuous; disc convex; base sinuous. Scutellar shield: Somewhat shiny, setose, punctures deep, close, subtriangular and distally rounded. Elytron: Convex, elongate, gradually narrowed from humeri to apices; conjoined tightly at apex; somewhat shiny, with elongate, yellow recumbent setae; length 2.0 mm, width 0.375 mm at humeri; humeri with punctate striae; disc with punctate striae; interstices slightly elevated, transversely rugose; apices without cavities. Legs: Femora and tibiae dark brown; tarsi dark reddish brown; surface somewhat shiny; punctures shallow, with yellow recumbent setae. Venter (Figure 38): Dark brown; surface somewhat shiny, with elongate, yellow recumbent setae; punctate, except along lateral side of hypomeron. Aedeagus (Figure 39): Elongate, dorsally flattened, sinuate, apically narrowed; median lobe free, short, apically rounded; parameres apically elongate, rounded, longer than median lobe, with lateral tooth; secondary lateral lobe absent; basal piece elongate, apically rounded. Female allotype (Figure 40): Length, 4.0 mm. Width, 1.0 mm. Antennae filiform, about 1/3 the length of the body; flagellomere I longer than wide, longer than II; flagellomere II–IV slightly longer than wide, sub-equal; flagellomeres V–VIII each slightly longer than IV, longer than wide, sub-equal. Antennae reddish-brown. Legs reddish brown. Lateral antennal grooves deeper, surfaces smooth, ridge medially undefined. Distribution. This new eucnemid species is known from two localities at La Selva (successional plot and primary forest) within Costa Rica. Biology. The holotype was taken from a successional plot. The allotype was taken from a primary forest. Larvae and pupae are unknown. Etymology. The specific epithet is derived from the dark brown coloration of the beetle., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 21-22

Adelothyreus totus Otto, Muona and Córdoba-Alfaro, new species Figures 17–19 Diagnosis. A complete lateral pronotal ridge will distinguish the new species from all known Adelothyreus species present in Costa Rica. Type material. Male holotype: “ COSTA RICA: Heredia pr.; La Selva Bio. Sta.; 3 km S Pto. Viejo; 10°26′N 84°01′W ” / “I. IV.1988; H.A. Hespenheide ” (“I”, “IV” and “8” handwritten) / “ HOLOTYPE:; Adelothyreus; totus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2021” (red printed label). Holotype to be deposited in INBC. Description. Male holotype: Length, 3.0 mm. Width, 0.75 mm. Body elongate; uniformly dark black; antennae dark brownish black; legs dark brown; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 17). Head: Surface densely punctate, somewhat dullish, subspherical; frons convex, without median fovea or carina; apical margin of frontoclypeal region rounded, more than 2.0 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Pectinate from flagellomeres III–VIII, attaining about 3/4 the length of the body; flagellomere I longer than II; flagellomere II strongly, asymmetrically serrate; flagellomere III longer than wide, shorter than IV; flagellomeres IV–VIII each sub-equal, longer than wide; flagellomere IX longer than VIII; rami on flagellomeres III–VIII arising at apical end, elongate and about 2.0 times longer than length of segment. Pronotum: Surface somewhat dullish, closely punctate; longer than wide, with poorly developed hind angles; lateral sides parallel-sided, apically arcuate; disc flattened with delicate median groove; anterior lateral pronotal ridge completely connected to the posterior lateral pronotal ridge forming a complete lateral ridge (Figure 18); base sinuous, strongly impressed above scutellar shield. Scutellar shield: Short, transverse, shallowly punctate, setose and distally truncated. Elytra: Shallow striate indicated at humeri, indistinct elsewhere; interstices flattened; surfaces shiny, closely punctate to rugose at basal 1/2, shallowly punctate at apical 1/2. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV very short, excavate; metatarsomere V short with simple claws. Venter (Figure 19): Closely punctate, with short, recumbent yellowish setae; hypomeron with apically widened notosternal antennal grooves; antennal grooves apically punctate, basally smooth; metepisterna caudally wide; elytral epipleura punctate; metacoxal plates parallel-sided. Distribution. This eucnemid species is only known from its type location in Costa Rica. Biology. No biological information is known for the new species. Larvae and pupae are unknown. Etymology. The specific epithet is derived from ‘totus’, a Latin adjectival word for complete due to the presence of a complete lateral pronotal ridge., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 9-10

Maelodrus costaricensis Otto, Muona and Córdoba-Alfaro, new species Figures 27–29 Diagnosis. This new eucnemid species is distinguished by its elongate bicolored form and vaguely defined lateral antennal grooves from all known eucnemid species present in Costa Rica. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “ FPM/23 /07; Pentaclethra macroloba; 14 Octubre 1994 ” / “ HOLOTYPE:; Maelodrus; costaricensis ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Description. Male holotype: Length, 3.0 mm. Width, 0.75 mm. Body elongate; bicolored with black head, dark brown pronotal disc and elytra, lateral pronotal sides, hypomera and abdomen infuscate reddish; antennae black; legs reddish-brown; head and elytra clothed with short, recumbent yellowish setae, pronotum with elongate, recumbent yellowish setae (Figures 27–28). Head: Surface evenly punctate, somewhat shiny, subspherical; frons convex, without median carina or fovea above frontoclypeal region; apical margin of frontoclypeal region rounded, about 2 times wider than base; interantennal carina absent; mandibles stout, bidentate, densely punctate. Antenna: Filiform to weakly serrate from flagellomeres I–IX, attaining about 3/4 the length of the body; flagellomere I longer than wide, slightly shorter than II; flagellomeres II–VIII each sub-equal, longer than wide; flagellomere IX simple, slightly longer than VIII. Pronotum: Surface somewhat dullish, densely punctate; slightly longer than wide, with moderate, sharp hind angles; lateral sides parallel-sided at basal 2/3, arcuate at apical 1/3; disc convex without fovea or circular fovea; base sinuous. Scutellar shield: Short, sub-triangular, setose, shallowly punctate and distally rounded. Elytra: Distinctly, shallowly striate; interstices slightly elevated; surfaces shiny with dense punctures. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV very short, excavated, slightly emarginated; metatarsomere V short with simple claws. Venter (Figure 29): Closely punctate, with elongate, recumbent yellowish setae; hypomeron with vaguely defined, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 6.0 times wider than laterally. Distribution. This eucnemid species is known from a holotype specimen taken from a single location within the province of Heredia. Biology. A single specimen was taken from Pracaxi (Pentacletha macroloba (Willdenow) Kuntze; Fabaceae). Larvae and pupae are unknown. Etymology. The specific epithet is derived from a combination of two words, ‘Costa Rica’ and ‘-ensis’, a Latin adjectival suffix meaning “pertaining to”; from which the new species have been taken., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 14-15

Dromaeolus americanus Otto, Muona and Córdoba-Alfaro, new species Figures 54–55 Diagnosis. This new eucnemid species can be readily distinguished from any ornate members of Dromaeolus present in Costa Rica by its elytral vittae and smaller size. The elytral vittae are very similar to an Amazonian eucnemid, D. batesii Bonvouloir, but D. americanus differs by its smaller size. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/06/198; 01 Septembre 1993; Bosque primario” / “ HOLOTYPE:; Dromaeolus; americanus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Paratype. 1 ♂: COSTA RICA: “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/01/320; 15 Enero 1994; Parcelas sucesionales” / “ PARATYPE:; Dromaeolus; americanus ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Paratype to be deposited in INBC. Description. Male holotype: Length, 4.5 mm. Width, 1.0 mm. Body elongate; black; antennae black; femur and tibiae black, tarsi black; head and pronotum clothed with short, recumbent silver-colored setae more apparent along base and lateral sides of pronotum, elytra with silver-colored setae more apparent with comma-shaped vittae at humeri, wide band across the apical 1/3 of each elytron, short, narrow patch along elytral suture just below scutellar shield (Figure 54). Head: Surface dullish, subspherical with closely spaced punctures; frons convex, simple with slight circular impression above interantennal carina; interantennal carina present; apical margin of frontoclypeal region weakly trilobed, more than 2 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Filiform from flagellomeres I–IX, attaining nearly 1/3 the length of the body, never passed beyond pronotal hind angles; flagellomere I longer than wide, longer than II; flagellomeres II–VIII each quadrate, sub-equal; flagellomere IX slightly longer than VIII. Pronotum: Surface dullish, with evenly dispersed punctures; slightly longer than wide, with well-developed, sharp hind angles; sides arcuate above pronotal hind angles; disc convex without shallow median groove or circular fovea; base sinuous, with delicate short median groove. Scutellar shield: Quadrate, sub-triangular, shallowly punctate and distally rounded. Elytra: Very shallow striae indicated; interstices slightly elevated; surfaces dullish, transversely rugose at basal 1/4, closely punctate elsewhere. Legs: First tarsomere longer than the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV excavated; metatarsomere V short with simple claws. Venter (Figure 55): Closely punctate, with short, recumbent silver-colored setae; hypomeron with basally opened, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Variation. The male paratype measured 5.0 mm long and 1.0 mm wide. It is slightly longer than and just as wide as the holotype. Interantennal carina is more apparent than the holotype. A circular impression above the interantennal carina is also more apparent than the holotype. Distribution. This species is known from two specimens taken from La Selva within the province of Heredia. Biology. The holotype was taken from a primary forest. The paratype was taken from a successional plot. Larvae and pupae are unknown. Etymology. The specific epithet is named after America, a name first used shortly after Christopher Columbus’ voyage to the new world, in which the name derives from Amerigo Vespucci, an Italian explorer who explored the new continents years later., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 24-25

Adelothyreus costaricensis Otto, Muona and Córdoba-Alfaro, new species Figures 13–16 Diagnosis. This new species is very similar to A. comes. The posteroventrally directed anterior lateral pronotal ridge will distinguish the new species from A. comes. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/01/332; 01 Febrero 1994; Parcelas sucesionales” / “ HOLOTYPE:; Adelothyreus; costaricensis ♂; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Female allotype: “ COSTA RICA, Heredia; Est.Bio. La Selva, 50–; 150 m, 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/15/121; 02 Junio 1993; Bosque secundario” / “ ALLOTYPE:; Adelothyreus; costaricensis ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Holotype and allotype are to be deposited in INBC. Description. Male holotype: Length, 2.5 mm. Width, 0.75 mm. Body elongate; uniformly dark black; antennae dark brownish black; femur and tibiae dark brownish black; tarsi dark brown; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 13). Head: Surface densely punctate, somewhat dullish, subspherical; frons convex, without median fovea or carina; apical margin of frontoclypeal region rounded, more than 2.0 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Pectinate from flagellomeres II–VIII, attaining about 2/3 the length of the body; flagellomere I longer than II; flagellomeres II–III sub-equal, each shorter than IV; flagellomere IV longer than III, shorter than V; flagellomeres V–VIII each sub-equal, longer than wide; flagellomere IX longer than VIII; ramus on flagellomere II arising along entire length, rami on flagellomeres III–VIII arising at apical end, elongate and about 1.5 times longer than length of segment. Pronotum: Surface somewhat dullish, closely punctate; longer than wide, with poorly developed hind angles; lateral sides parallel-sided, apically arcuate; disc convex without median groove; anterior lateral pronotal ridge short, arcuate, directed posteroventrally; posterior lateral pronotal ridge elongate, about 3/4 the length of the pronotum (Figure 14); base sinuous, with circular strong impressions above scutellar shield. Scutellar shield: Short, quadrate, sub-triangular, shallowly punctate, setose and distally rounded. Elytra: Very shallow striate indicated; interstices slightly elevated; surfaces shiny, very closely punctate to rugose at basal 1/2, closely punctate at apical 1/2. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV very short, excavate; metatarsomere V short with simple claws. Venter (Figure 15): Closely punctate, with short, recumbent yellowish setae; hypomeron with apically widened notosternal antennal grooves; antennal grooves smooth, sparsely punctate; metepisterna caudally wide; elytral epipleura punctate; metacoxal plates parallel-sided. Female allotype (Figure 16): Length, 3.0 mm. Width, 1.0 mm. Antennae serriform, about 1/2 the length of the body; flagellomere I longer than wide, longer than II; flagellomere II quadrate, serrate, shorter than III; flagellomeres III–VIII sub-equal, transverse, asymmetrically serrate; flagellomere IX longer than VIII. Distribution. This species is only known from its type location in Costa Rica. Biology. The holotype was taken from a successional plot of trees. The allotype was taken from a secondary growth forest. Larvae and pupae are unknown. Etymology. The specific epithet is named for the country of Costa Rica from which the new species was taken., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 8-9

More than 1300 specimens of Eucnemidae collected from Heredia Province in Costa Rica during the1990s Arthropods of La Selva (ALAS) survey were studied from 2018 through 2022. One new genus of falseclick beetle, Absensiugum Otto, Muona and Córdoba-Alfaro, is described. Nematodes teres Horn, from the Ne-arctic and Caribbean regions, is transferred to this new genus to form Absensiugum teres, new combination.Sixteen new species of false click beetle (Coleoptera: Eucnemidae) are described from Costa Rica. These newspecies are: Adelothyreus brevis, Adelothyreus costaricensis, Adelothyreus totus, Quirsfeldia stethonoides, Lacuspectinatus, Maelodrus costaricensis, Onichodon confluentus, Onichodon rufus, Isarthrus striatus, Absensiugumbrunneum, Dromaeolus americanus, Dromaeolus brunneus, Dromaeolus herediensis, Dromaeolus holdridgei,Deltometopus bicolor and Nematodes apicalis. Three additional records outside of the Heredia Province fromthe Osa Peninsula and Panama for Lacus pectinatus are included in this study. Identification keys are providedfor species of Adelothyreus Chevrolat, Onichodon Newman, Dromaeolus Kiesenwetter, Deltometopus Bonvou-loir and Nematodes Berthold in Costa Rica. Diagnostic differences are briefly noted for each species within theNeotropical region. A list of Eucnemidae from Heredia Province is provided.

Quirsfeldia stethonoides Otto, Muona and Córdoba-Alfaro, new species Figures 20–22 Diagnosis. Generic character features like caudally obliterated notosternal antennal grooves, parallel-sided metacoxal plates and apical margin of the frontoclypeal region being less than 2.0 wider than the base will distinguish the new species from all known species within the tribe in Costa Rica. Type material. Female holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/06/170; 03 Agosto 1993; Bosque primario” / “ HOLOTYPE: Quirsfeldia; stethonoides ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (red printed label). Holotype to be deposited in INBC. Description. Female holotype: Length, 3.0 mm. Width, 1.0 mm. Body elongate; uniformly dark brownish black; antennae dark brownish black; legs brown; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 20). Head: Surface densely punctate, somewhat dullish, subspherical; frons convex, with delicate median fovea above frontoclypeal region; apical margin of frontoclypeal region rounded, less than 2.0 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Asymmetrically serrate from flagellomeres II– IX, presumably attaining about 1/3the length of the body, never beyond pronotal hind angles; flagellomere I longer than II; flagellomere II shorter than I and III, quadrate; flagellomeres III–VII each sub-equal, wider than long; flagellomeres VIII–IX missing on right antenna; flagellomeres VI–IX missing on left antenna. Pronotum: Surface somewhat dullish, granulose; slightly longer than wide, with moderate, sharp hind angles; lateral sides parallel-sided at basal 2/3, arcuate at apical 1/3; disc convex with short, delicate median groove above scutellar shield; anterior lateral pronotal ridge short, arched, ventrally bent; posterior lateral pronotal ridge elongate, extending almost to caudal end (Figure 21); base sinuous. Scutellar shield: Short, sub-triangular, shallowly punctate, with median groove and distally rounded. Elytra: Shallow striate indicated; interstices slightly elevated; surfaces shiny, transversely rugose at basal 1/2, dense, rugose at apical 1/2. Legs: First tarsomere shorter than the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–IV simple; metatarsomere IV very short; metatarsomere V elongate with simple claws. Venter (Figure 22): Closely punctate, with short, recumbent yellowish setae; hypomeron with caudally obliterated, poorly defined notosternal antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates parallel-sided. Distribution. This eucnemid species is only known from its type location in Costa Rica. Biology. The holotype was taken from a primary forest. Larvae and pupae are unknown. Etymology. The specific epithet is derived from its similar appearance to the Eucnemine genus, Stethon LeConte. Subfamily Macraulacinae Fleutiaux, 1923 Diagnosis. Form oblong, elongate or obtuse; antennomeres usually sexually dimorphic; mandibles either stout with a basal tooth or slender without teeth; simple lateral pronotal ridge present; hypomeron either simple, with basally closed lateral antennal grooves or with basally open lateral antennal grooves; legs slender; prothoracic tibiae with one apical spur; lateral surfaces of mesothoracic and metathoracic tibiae usually with transverse rows of spines; tarsomere IV often bilobed; tarsal claws either simple or basally toothed; prothoracic tarsomere I usually with basal sex combs in males; male aedeagus with dorsally open basal piece; median lobe simple, with solidly fused slender basal struts; fused to lateral lobes; lateral lobes entire, either with notched or apically deeply and narrowly bifurcate; bursa either simple or divided; spermatheca tripartite, sclerotized, divided (Muona 1993; Otto 2016). Tribe Macraulacini Fleutiaux, 1923 Diagnosis. Form oblong, elongate or obtuse; flagellomeres usually sexually dimorphic; mandibles either stout with a basal tooth or slender without teeth; simple lateral pronotal ridge present; hypomeron either simple, with basally closed lateral antennal grooves or with basally open lateral antennal grooves; legs slender; prothoracic tibiae with one apical spur; lateral surfaces of mesothoracic and metathoracic tibiae usually with transverse rows of spines; tarsomere IV often bilobed; tarsal claws either simple or basally toothed; prothoracic tarsomere I usually with basal sex combs in males; male aedeagus with dorsally open basal piece; median lobe simple, with solidly fused slender basal struts, fused to lateral lobes; lateral lobes entire, either with notched or apically deeply and narrowly bifurcate; bursa either simple or divided; spermatheca tripartite, sclerotized, divided (Muona 1993; Otto 2017c)., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on page 11, {"references":["Muona J. 1993. Review of the phylogeny, classification and biology of the family Eucnemidae (Coleoptera). Entomologica Scandinavica Supplement 44: 1 - 133.","Otto RL. 2016. The false click beetles (Coleoptera: Eucnemidae) of Laos. Entomologica Basiliensia et Collectionis Frey 35: 181 - 427.","Otto RL. 2017 c. Descriptions of six new species of false click beetles (Coleoptera: Eucnemidae: Macraulacinae) with new identification keys for one tribe and two genera. Insecta Mundi 0558: 1 - 19."]}

Absensiugum Otto, Muona and Córdoba-Alfaro, new genus Type species. Absensiugum brunneum new species, by present designation. Diversity and distribution. Absensiugum new genus is a genus consisting of two species distributed in the Caribbean, Nearctic and Neotropical regions. The new species was found in two locations within a single area in Costa Rica. The second species is distributed in the Caribbean region and southeastern United States. Description. Male. Body elongate, approximately 3.5 times longer than wide, dorsally convex and ventrally well sclerotized. Head: Hypognathous, with short, recumbent setae. Antennae tubular, filiform with 11 antennomeres, setose; scape 3.0 times longer than pedicel; pedicel globular, shorter than flagellomere I; flagellomere I longer than II; flagellomeres II–IV sub-equal in length and rounded in cross section; flagellomeres V–VIII sub-equal in length and rounded in cross section; flagellomere IX longer than VIII. Eyes round, well developed, enlarged, not incised. Frontoclypeal region subtriangular, apically trilobed, about 2.5 times wider than base. Mandibles well developed, stout, bidentate. Maxillary and labial palpi concealed behind mandibles. Labrum concealed. Pronotum: Subparallel-sided, convex and setose. Longer than wide, with moderately sized pronotal hind angles. Lateral pronotal ridge entire, straight. Disc convex. Base sinuous. Scutellar shield: Setose, short, sub-triangular. Elytron: Elongate, convex, setose. Disc with striae indicated as punctate lines. Interstices slightly elevated. Cavities absent near elytral apices. Legs: Prothoracic legs shortest, metathoracic legs longest. Protibia apically rounded, setose, with one apical spur. Lateral sides of meso- and metatibiae with setae and transverse row of spine combs. Metatarsi, including claws, as long as tibia. Protarsomere I with straight, basal sex combs. First metatarsi as long as combined length of remaining four. Metatarsi I–III simple. Metatarsus IV excavate-emarginate, as wide as III. Metatarsus V elongate. Pretarsal claws basally toothed. Tarsal formula 5-5-5. Venter: Surface with recumbent setae. Prosternal peg basally narrow, elongate. Notosternal suture as long as the hypomeral base. Hypomeron without medially defined antennal ridge, surface smooth along lateral ridge, rest punctate. Epipleura simple, not grooved. Metepisterna parallel-sided. Metacoxal plate medially 3.0–6.0 times wider than laterally. Tarsal grooves absent on meso- and metasterna. Abdomen with 5 visible ventrites, convex medially. Last visible ventrite rounded caudally. Etymology. The new generic name is a combination of two Latinized words, absens, which means absent and iugum which means ridge, due to the missing medial antennal ridge of the hypomeron. Gender neuter. Note. During the course of the study, R. Otto discovered a pair of Costa Rican beetles superficially resembling the Nearctic species, Dromaeolus teres (Horn). Upon further examination, these two species were found to share the same morphological attributes, resulting in our decision to create a new group name, Absensiugum new genus, for these species. Therefore, we propose to transfer Nematodes teres Horn to the new group, hereby recognized as Absensiugum teres (Horn) new combination., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on page 20

Onichodon confluentus Otto, Muona and Córdoba-Alfaro, new species Figures 30–32 Diagnosis. The new eucnemid species can be readily distinguished by the presence of confluent surfaces of the frons from any members of Fornax and Onichodon present in Costa Rica. Type material. Male holotype: “ COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/15/121; 02 Junio 1993; Bosque secundario” / “ HOLOTYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro’ Det. R. L. Otto; 2019” (red printed label). Female allotype: “ COSTA RICA: Prov. Heredia:; 16 km SSE La Virgen, 1050–; 1150 m, 10°16′N 84°05′W; 9 Abril 2001; INBio-OET-ALAS transect” (“9 Abril”, “1” were handwritten on label) / “ 09 Abril 2001; 11/M/01/061; Transect” / “ ALLOTYPE:; Onichodon; confluentus ♀; Otto, Muona & Córdoba-Alfaro; Det. R. L. Otto; 2019” (yellow printed label). Holotype and allotype are to be deposited in INBC. Paratypes. 7 ♂♂: COSTA RICA: 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/14/061; 02 Abril 1993; Bosque secundario” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/15/094; 03 Mayo 1993; Bosque secundario” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/16/122; 02 Junio 1993; Bosque secundario” / “Collection of the Global; Eucnemid Research Project; (Robert L. Otto)” (green framed white label) / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (GERP); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/14/148; 01 Julio 1993; Bosque secundario” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/12/397; 30 Junio 1995; Bosque primario” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “COSTA RICA: Heredia; Est.Bio. La Selva, 50–; 150 m 10°26′N 84°01′W; Proy. ALAS, INBio-OET” / “M/05/655; 31 Mayo 1996; Bosque primario” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC); 1 ♂, “ 21 Mar 2001; INBio-OET-ALAS transect; COSTA RICA: Prov. Heredia:; 16 km SSE La Virgen, 1050–; 1150 m, 10°16′N 84°05′W (“21 Mar” and “1” were handwritten on label) / 21 Marzo 2001; Transect; 11/M/07/067” / “ PARATYPE:; Onichodon; confluentus ♂; Otto, Muona & Córdoba-Alfaro; Det. R.L. Otto; 2019” (yellow printed label) (INBC). Paratypes are to be deposited in INBC and GERP. Description. Male holotype: Length, 8.0 mm. Width, 2.5 mm. Body elongate; dark reddish-brown; antennae and legs dark reddish-brown; head, pronotum and elytra clothed with short, recumbent yellowish setae (Figure 30). Head: Surface dullish, subspherical, confluently rugose; frons convex, simple without impression or carina; apical margin of frontoclypeal region weakly trilobed, about 2 times wider than base; mandibles stout, bidentate, densely punctate. Antenna: Filiform from flagellomeres I–IX, attaining at least 1/3 the length of the body, just beyond pronotal hind angles; flagellomere I longer than wide, slightly longer than II; flagellomeres II–VIII each longer than wide, sub-equal; flagellomere IX slightly longer than VIII. Pronotum: Surface somewhat shiny, with closely spaced punctures; quadrate, with well-developed, sharp hind angles; sides subparallel-sided at basal 1/2, arcuate at apical 1/2, slightly sinuate above pronotal hind angles; disc convex with shallow fovea above scutellar shield; base sinuous, with circular pit on each side above scutellar shield. Scutellar shield: Longer than wide, sub-triangular, shallowly punctate, setose and distally rounded. Elytra: Very shallow striae indicated; interstices slightly elevated; surfaces shiny, transversely rugose at basal 1/4, closely punctate elsewhere. Legs: First tarsomere as long as the combined lengths of the remaining four on meso- and metatarsi; tibiae rounded in cross section; metatarsomeres I–III simple; metatarsomere IV excavated; metatarsomere V elongate with basally toothed claws. Venter (Figure 31): Closely punctate, with short, recumbent yellowish setae; hypomeron with basally opened, lateral antennal grooves; metepisterna parallel-sided; elytral epipleura punctate; metacoxal plates medially 3.0–6.0 times wider than laterally. Female allotype (Figure 32): Length, 12.0 mm. Width, 4.0 mm. Antennae weakly and asymmetrically serriform, about 1/3 the length of the body; flagellomere I longer than wide, slightly longer than II; flagellomere II slightly longer than wide, shorter than III; flagellomeres III–VIII longer than wide, sub-equal. Variations. Seven male paratypes measured 6.0– 10.5 mm long and 1.5–2.5 mm wide. Two male paratypes are larger than and just as wide as the holotype. The remaining five paratypes are shorter and narrower than the holotype. Pronotal punctures are widely dispersed in one paratype, whereas the remaining six paratypes have exoskeletal structures similar to the holotype. Distribution. This eucnemid species is known from nine specimens taken from two locations within the province of Heredia. Biology. Two specimens were taken from a primary forest. Five specimens, including the holotype were taken from a second growth forest. Larvae and pupae are unknown. Etymology. The specific epithet is named from the presence of confluent surfaces of the frons on the new species., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on pages 16-17

Genus Maelodrus Fleutiaux, 1928 Diversity and distribution. Five described species and numerous undescribed species are known in the Oceanic region from the Philippines and Indonesia to Samoa. The new Central American species is present within a single province in Costa Rica. A second, undescribed species is present in Peru (Vahtera et al. 2015). Diagnosis. Apical margin of frontoclypeal region fairly evenly rounded and more than twice as wide as the distance between antennal sockets; vaguely defined, either shallow or deep, basally opened lateral antennal grooves present; male protarsomere I simple, with straight, basal sex combs; elytral apices meeting tightly together; metacoxal plates medially at least 6.0 times wider than laterally; frons simple; last visible ventrite either rounded, emarginated or acute; simple tarsal claws; lateral surfaces of meso- and metatibiae either with setae and transverse rows of spine combs or setae and irregularly placed flat spines (Muona 2011). These diagnostic characteristics, especially the presence of a vaguely defined lateral antennal groove and elongate form will distinguish the group from any genera within the tribe Macraulacini in Costa Rica., Published as part of Otto, Robert L., Muona, Jyrki & Córdoba-Alfaro, Jim, 2023, A new genus and sixteen new species of false click beetles (Coleoptera: Eucnemidae) described from the Heredia Province of Costa Rica with several additional records from the Osa Peninsula and Panama, pp. 1-36 in Insecta Mundi 2023 (991) on page 14, {"references":["Vahtera V, Muona J, Linna A, Saaksjarvi I. 2015. Nine genera of Eucnemidae (Coleoptera) new to Peru, with key to Peruvian genera. Biodiversity Data Journal 3: e 4493. doi: 10.3897 / BDJ. 3. e 4493.","Muona J. 2011. Eucnemidae. info Homepage. Available at http: // dol. luomus. fi: 8080 / cgibin / dol / dol _ homepage. pl (Last accessed 15 August 2019.)"]}

Asiocnemis colombicus Muona, new species Figures 8–11. Etymology. Named after the Republic of Colombia. Holotype. Embedded in a flat, quadratic Colombian resin piece, 55 mm × 30 mm, 4 mm thick, female. Diagnosis. The elongated, parallel-sided and flat shape, wide smooth frontoclypeal area, slender tarsi with tarsomere 4 feebly bilobed, simple, minute claw and slender antennae separate this species from all previously known neotropical Dromaeolus species as well as all American Asiocnemis Mamaev species. Description. Length 5.5 mm. Elateroidea synapomorphy: pro-mesocoxal clicking mechanism present. Eucnemidae synapomorphies: pedicel attached subapically to scape, labrum hidden, elytral striae with apical excretory punctures. Macraulacinae synapomorphy: hypomera with basally open lateral antennal grooves (Fig. 9). Macraulacini diagnostic other characters: tibiae with lateral spine-combs. Form narrow, parallel sided, flat (Fig. 8). Head rounded, densely punctate, eyes medium-sized, frontoclypeal region wide, 1.3 times as wide apically as between antennal sockets, apical margin slightly rounded, mandibles short, bifid. Pronotum elongated, about as wide as long, with short, deep basal median groove converging craniad and ending before middle, with acute non-carinate hind angles, roughly and very densely punctate, scutellum strongly transverse, fairly large, parallel-sided, slightly bilobed caudally, elytra densely and roughly punctate with strong sutural striae and apical excretory punctures, disk without striae. Antennal sockets round, antennae moderate long, slender, f8–f9 reaching past pronotal hind angles, scape long, elongate, pedicel shorter than f1, all flagellomeres longer than wide, becoming more elongated towards apex, f4 twice as long as wide, f9 longest of all, 1.5 times longer than f8 (Fig. 9, 10). Hypomeron densely punctate with sharply delimited non-punctate lateral antennal grooves (Fig. 9, 10), prosternal peg parallel-sided, apically pointed, prosternum very densely punctate, metasternum very densely and roughly punctate. Metacoxal plates triangular, very densely punctate, widest close to midline, three times as wide as on sides, less so at midline, caudal angle rounded. Legs long, elongate, protarsi shorter than protibiae, tarsomeres 2–4 increasingly narrower and tarsomere 5 small, delicate, meso- and metatibiae long, slender, longer than tarsi, with strong spine-combs on lateral surfaces, meso- and metatarsomere 1 as long 2–4 combined, 4 excavated, slightly wider than previous one, claws small, simple (Fig. 11). Abdominal ventrites connate, very densely punctate. Remarks. As pointed out by Muona (2000), Dromaeolus is an artificial assemblage of species waiting for a phylogenetic analysis. Such an attempt requires the inclusion of all Macraulacini genera, a gargantuan task. The present species does not fit any previously known Dromaeolus in the only “complete” key available (Bonvouloir 1871), nor does it fit any species described from South America. It is also clear that it is not congeneric with the European Dromaeolus barnabita Villa, the type-species of this genus (Muona 1987). The genus Asiocnemis differs from Dromaeolus by conspicuous larval characters but it has not been satisfactorily defined with external adult characters. The present species with slender, parallel form, delicate tarsi, basally strongly grooved elongate pronotum and rough surface sculpture fits the known species well and it has been placed in Asiocnemis. There are seven previously reported Asiocnemis species from the Americas, distributed in the western mountains from Washington to Ecuador and French Guiana. It is assumed that several South American slender black Dromaeolus species belong here as well. Whether this newly described species is a recent one or an extinct creature from the past, cannot be determined at this time. A somewhat similar one, Asiocnemis sp., was reported from Baltic amber (Muona 1993a)., Published as part of Muona, Jyrki, 2022, New fossil false click-beetles from the Americas (Coleoptera: Eucnemidae), pp. 1-11 in Insecta Mundi 2022 (912) on pages 9-10, DOI: 10.5281/zenodo.6391857, {"references":["Muona J. 2000. A revision of the North American Eucnemidae. Acta Zoologica Fennica 212: 1 - 106.","Bonvouloir HA de. 1871. Monographie de la Famille des Eucnemides, 1 st part. Annales de la Societe entomologique de France 40 (Supplement) 1 - 288 + 21 pls.","Muona J. 1987. The generic names of the beetle family Eucnemidae (Coleoptera). Entomologica Scandinavica 18: 79 - 92.","Muona J. 1993 a. Eucnemidae and Throscidae from Baltic amber (Coleoptera). Entomologische Blatter fur Biologie und Systematik der Kafer 89: 15 - 45."]}

Thambus woodruffi Muona, new species Figures 6, 7. Etymology. Named in honor of the late Robert E. Woodruff, whose many interests included fossil resins and especially the Dominican amber fauna. Holotype. Embedded in flat, rectangular Dominican resin piece with broadly rounder corners, 17 mm × 11 mm, female. The same piece includes an unidentified throscid beetle as well as other insects (Fig. 7). Diagnosis. The length/width ratio of all other Thambus species is 2.4–2.8, in this species it is about 2.1. Description. Elateroidea synapomorphy: pro-mesocoxal clicking mechanism present. Eucnemidae synapomorphies: pedicel attached subapically to scape, labrum hidden, elytral striae apically grooved with excretory punctures (Fig. 6). Eucnemidae other characters: abdominal ventrites connate (Fig. 7). Macraulacinae synapomorphy: hypomera with basally open lateral antennal grooves (Fig. 7). Macraulacini other characters: hypomera without excretory pits, metasternum without tarsal grooves. Length 3.0 mm. Form very wide, ratio L/W about 2.1 (Fig. 6). Pronotum wider than long and wider than elytra with slightly expanded, acute non-carinate hind angles, evenly and densely punctate (Fig. 6). Scutellum transverse, widely triangular, apex cut off, elytra with sharp, complete sutural striae with deep apical excretory groove continuing around to lateralmost striae, densely and finely punctate, other striae faint but present (Fig. 6). Head rounded, densely punctate, eyes large, flat, frons wide, without carinae, frontoclypeus apically about 1.3 times as wide as between antennal sockets, mandibles short, bifid (Fig. 7). Antennae fairly short, reaching slightly past the hind angles, scape as long as pedicel plus f1 and f2 combined, f2–f3 feebly serrate, f4–f7 not visible, f9 1.4 times as long as f8, both longer than wide. Hypomeron with sharply defined, basally open lateral antennal grooves, about triangular, sparsely punctate, without excretory pits (Fig. 7). Prosternum wide, moderately densely punctate, prosternal peg long, triangular, acute, deeply marginate, metasternum wide and long, densely punctate, without tarsal grooves, elytral epipleura simple. Metacoxal plates triangular, widest close to midline, angle tightly rounded and there less than times as wide as on sides, densely punctate. Legs long and delicate, meso- and metatarsi shorter than tibiae, meso- and metatarsomere 1 slightly shorter than the others combined, fourth apically narrowly bilobed, excavated, claw simple (Fig. 6, 7). Abdominal ventrites connate, densely and finely punctate (Fig. 7). Remarks. Thambus Bonvouloir is a primarily tropical American genus, with additional species known from Europe (1), China (1), Japan (1) and North America (1). The species are small, have a basally wide pronotum with sides strongly converging craniad. In addition to their shape and small size, Thambus species are characterized by round pit-like areas laterally on the prosternum. This feature varies between species from faint (e.g. T. deyrollei Bonvouloir) to conspicuous (e.g. T. friwaldskyi Bonvouloir). These pits are not visible in the present sample, but the other characters listed place it to Macraulacini and within this tribe the correct placement is clearly Thambus. Family Eucnemidae Eschscholtz, 1829 Subfamily Macraulacinae Fleutiaux, 1922 Tribe Macraulacini Fleutiaux, 1922, Published as part of Muona, Jyrki, 2022, New fossil false click-beetles from the Americas (Coleoptera: Eucnemidae), pp. 1-11 in Insecta Mundi 2022 (912) on pages 6-9, DOI: 10.5281/zenodo.6391857

Muona, Jyrki (2022): New fossil false click-beetles from the Americas (Coleoptera: Eucnemidae). Insecta Mundi 2022 (912): 1-11, DOI: http://doi.org/10.5281/zenodo.6391857, {"references":["Bonvouloir HA de. 1871. Monographie de la Famille des Eucnemides, 1st part. Annales de la Societe entomologique de France 40(Supplement) 1-288 + 21 pls.","Carter JG, Gallagher PE, Valone RE, Rossbach TJ. 1988. Fossil collecting in North Carolina. Bulletin 89. Department of Natural Resources and Community Development; Raleigh, NC. 86 p.","Chang H, Muona J, Hanyong P, Li, X, Chen W, Teravainen M, Dong R, Qiang Y, Xingliao Z, Songhai J. 2016. Chinese Cretaceous larva exposes a southern Californian living fossil (Insecta, Coleoptera, Eucnemidae). Cladistics 32: 211-214.","Kosmowska-Ceranowicz B. 1999. Succinite and some other fossil resins in Poland and Europe (deposits, finds, features and differences in IRS). Estudios del Museo de Ciencias Naturales de Alava 14: 73-117.","Lawrence JF, Muona J, Teravainen M, Stahls G, Vahtera V. 2007. Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). Insect Systematics & Evolution 38: 205-239.","Lawrence JF, Slipinski A. 2013. Australian beetles. Volume 1, morphology, classification and keys. CSIRO Publishing; Collingwood, Australia. 566 p.","Li H, Chang H, Muona J, Zhao Y, Ren D. 2021. Subfamily Anischiinae (Coleoptera: Eucnemidae) in early Cretaceous of Northeast China. Insects 12: 105.","Li YD, Tihelka E, Liu ZH, Huan D, Cai CJ. 2020. Muonabuntor gen. nov., a new genus of false click beetles from mid- Cretaceous Burmese amber (Coleoptera: Elateroidea: Eucnemidae). Paleoentomology 003(4): 399-406.","Mickle JE. 1996. Grexlupus carolinensis, a new probable lauraceous fruit from the late Cretaceous of North Carolina. Journal of the Elisha Mitchell Scientific Society 112: 1-6.","Muona J. 1987. The generic names of the beetle family Eucnemidae (Coleoptera). Entomologica Scandinavica 18: 79-92.","Muona J. 1993a. Eucnemidae and Throscidae from Baltic amber (Coleoptera). Entomologische Blatter fur Biologie und Systematik der Kafer 89: 15-45.","Muona J. 1993b. Review of the phylogeny, classification and biology of the family Eucnemidae (Coleoptera). Entomologica Scandinavica Supplement 44: 1-133.","Muona J. 2000. A revision of the North American Eucnemidae. Acta Zoologica Fennica 212: 1-106.","Muona J. 2020. The late Cretaceous amber Eucnemidae fossils from Myanmar. Entomologische Blatter fur Biologie und Systematik der Kafer 116: 27-55.","Muona J. 2021. False click-beetles from Baltic amber (Coleoptera, Eucnemidae). Entomologische Blatter fur Biologie und Systematik der Kafer 117: 90-113.","Muona J, Chang H, Ren D. 2020. The clicking Elateroidea from Chinese Mesozoic deposits (Insecta, Coleoptera). Insects 11: 875.","Muona J, Teravainen M. 2020. A re-evaluation of the Eucnemidae larval characters (Coleoptera). Papeis Avulsos de Zoologia; v.60.special-issue: e202060 (s.i.).28.","Oberprieler R, Ashman L, Friese M, Slipinski A. 2016.The first elateroid beetles (Coleoptera: Polyphaga: Elateroidea) from the Upper Jurassic of Australia. Zootaxa 4147: 177-191.","Otto RL. 2019. Descriptions of two new elateroid beetles (Coleoptera: Eucnemidae, Elateridae) from Burmese amber. Insecta Mundi 702: 1-6.","Poinar G. 2012. Discoclavata dominicana n. gen., n. sp., (Coleoptera: Bostrichidae) and Lissantauga epicrana n. gen., n. sp. (Coleoptera: Eucnemidae) in Dominican amber. Historical Biology: An International Journal of Paleobiology 25: 107-113.","Ragazzi E, Roghi G, Giaretta A, Gianolla P. 2003. Classification of amber based on thermal analysis. Thermochimica Acta 404: 43-54.","Wickham HF. 1916. The fossil Elateridae of Florissant. Bulletin Museum Comparative Zoology 60: 493-527 + 7 pls.","Woodruff R. 2004. Amber versus Copal. Available at http://www.fossilmall.com/Science/AmberCopal.htm (Last accessed January 2022.)"]}

The phylogeny of the genus Arrhipis Bonvouloir (Coleoptera, Eucnemidae) is clarified with a cladistic analysis based on five molecular markers and morphology. Sixteen species from Africa, America, Asia, and Australia are included in the analysis. Two separate Asian clades are recovered, one of them being the sister group to a clade with the American and African species. With the exception of the continental south-east Asian species, all Gondwanan regions have monophyletic faunas. According to the present data, the continental south-east Asian fauna comprises two monophyletic groups, one of which is the sister group to African and American species. Vicariance seems to be the logical explanation for the distribution of these lignicolous beetles. © The Willi Hennig Society 2009.

Three eucnemid genera and species are recognized for the first time from Korea: Eutyptycluts vicinus Fleutiaux, 1923, Fornax consobrinus Hisamatsu, 1963, and Feaia nipparensis (Hisamatsu, 1957) new combination (= Heterotaxis nipparensis Hisamatsu, 1957).A key to species, diagnoses, redescriptions, and photographs of diagnostic characters are provided for known Korean species. (C) 2017 Published by Elsevier B.V. on behalf of Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society.

The larval characters of Eucnemidae are re-evaluated. Tribe Schizophilini Muona, 1993 is considered to merit subfamily rank as Schizophilinae Muona new status. Larval characters congruent with previously used adult morphological characters support the earlier published sister-group hypotheses for lignicolous eucnemids with one exception, Pseudomeninae, which is here split resulting in the following evolutionary hypothesis: (Pseudomeninae (Schizophilinae (Paleoxeninae (Melasinae, Eucneminae, Macraulcinae)))). The position of three remaining clades, Anischiinae, Perothopinae and Phyllocerinae remain ambiguous.

The genusDirrhagofarsusis firstly recorded from Korea with three species:Dirrhagofarsuslewisi(Fleutiaux, 1900),Dirrhagofarsusmodestus(Fleutiaux, 1923), andDirrhagofarsusunicolor(Hisamatsu, 1960). A key to Korean species ofDirrhagofarsus, with diagnoses, redescriptions, and photographs of important structures is provided. In this work, Dirrhagusmodestusf.unicolor Hisamatsu, 1960 is regarded as a valid species,Dirrhagofarsusunicolor(Hisamatsu, 1960),comb. n.

The genus Dirrhagofarsus is firstly recorded from Korea with three species: Dirrhagofarsus lewisi (Fleutiaux, 1900), Dirrhagofarsus modestus (Fleutiaux, 1923), and Dirrhagofarsus unicolor (Hisamatsu, 1960). A key to Korean species of Dirrhagofarsus, with diagnoses, redescriptions, and photographs of important structures is provided. In this work, Dirrhagus modestus f. unicolor Hisamatsu, 1960 is regarded as a valid species, Dirrhagofarsus unicolor (Hisamatsu, 1960), comb. n.

Palaeoxenus sinensis Chang, Muona & Teravainen sp. nov. (Coleoptera, Eucnemidae) is described on the basis of a Cretaceous larva found from the Yixian Formation in Huangbanjigou, Liaoning Province, China. The only previously known member of this clade is a southern Californian endemic, Dohrn's elegant eucnemid beetle (Palaeoxenus dohrni), a species that develops in conifers, especially the incense cedar (Calocedrus decurrens). The new find proves that the highly specialized main eucnemid lineages had evolved 123 Mya, before the main radiation of the angiosperms and probably as an adaptation to development in gymnosperms.

Thirteen genera of Eucnemidae containing forty species were collected from the Iquitos region in Peru. Nine of the genera are new to the country: Rhagomicrus Fleutiaux, 1902, Adelorhagus Horn, 1890, Adelothyreus Chevrolat, 1867, Microrhagus Dejean, 1833, Dyscharachthis Blackburn, 1900, Heterotaxis Bonvouloir, 1871, Spinifornax Fleutiaux, 1926, Serrifornax Fleutiaux, 1926 and Maelodrus Fleutiaux, 1928. The previous eucnemid record from Peru contained eleven species in ten genera. Only one of the forty species caught, Entomophthalmus americanus Bonvouloir, was previously known and described from the country. Dyscharachthis, Maelodrus and Adelorhagus are recorded from South America for the first time. Many of the collected species seem to favor white-sand forest as their habitat. Possible reasons for this are discussed. A list of eucnemids from Peru is included, containing taxa already recorded from the country and also taxa that are likely to occur there. A key to the Peruvian genera is included.

Recent findings in the greater Helsinki region of Hylochares cruentatus (Coleoptera, Eucnemidae), previously considered locally extinct, have shown ecological and morphological differentiation from H. cruentatus found in Russian Karelia. Despite their identical genetic constitution, differences in adult morphology, male genitalia as well as greatly differing life-history traits between the Finnish and Russian beetles prompted recognizing the Russian Hylochares as a distinct, new species: Hylochares populi n.sp., type locality Russia, Karelia, Kuganavalok. Single old specimens from Estonia and the Altai Mountains belong to the same taxon. Traditional morphological characters and life-history traits separate the two taxa, whereas the four applied genetic markers were identical, suggesting very recent diversification. The Russian H. populi lives in large soft-wooded aspen trees; H. cruentatus prefers tree- and bush-sized willows with harder wood. Conservation actions have to change focus from aspen trees to willow habitats to secure the survival of this rare beetle species in Finland. Zusammenfassung Die in der weiteren Umgebung Helsinkis in Finnland kurzlich gefundenen Hylochares cruentatus (Coleoptera, Eucnemidae), sie galten dort als lokal ausgestorben, weisen okologische sowie morphologische Differenzen zu dem im russischen Karelien vorkommenden H. cruentatus auf. Trotz des identischen genetischen Erbguts differieren die mannlichen Genitalien beider Kafer, zudem zeigen sie hochst verschiedene Lebensweisen. Aus diesen Grunden sollten russische Hylochares als eigenstandige Art eingeordnet werden: Hylochares populi n.sp. Typus-Lokalitat Russland, Karelien, Kuganavalok. Einzelne altere Exemplare aus Estland und dem Altaigebirge gehoren offenkundig zu dem gleichen Taxon wie auch die russisch-karelischen Kafer. Die beiden Taxa sind aufgrund traditioneller morphologischer Merkmale und Abweichungen der Lebensweise zu unterscheiden. Da sich hier die vier analysierten, genetischen Marker jedoch als identisch erweisen, lasst dies auf eine erst kurzlich stattgefundene Diversifizierung schliesen. Wahrend die russischen H. populi in grosen, weichholzigen Espen leben, bevorzugen H. cruentatus kleinere Weiden mit harterem Holz. Artenschutzmasnahmen in Finnland sollen sich daher nicht auf Espengebiete, sondern auf die Sicherung von Weidenbestanden konzentrieren, um das Uberleben dieses auserst seltenen Kafers zu sichern.

The function of microtrichial patches and areoles found in most eucnemid larvae is discussed. The areoles are regarded to be water excreting organs important for larval water-balance. The microtrichial patches were known to be an essential part of the larval locomotory method. It is suggested the modified spiracular collars associated with larvae of certain Dirhagini and Macraulacinae play a similar role. This appears especially clear with respect to Asiocnemis spp. The overall similarity between soft-bodied larvae of the few true wood-boring Eucnemidae is considered a homoplastic adaptation to multiple secondary invasions into relatively hard wood.

The biology of Hylochares cruentatus in Southern Finland is discussed. Contrary to previous observations in Russian Karelia, the species breeds in barkless willows along a regularly flooding small river in Southern Finland. The specialized biological requirements suggest this species might well survive in similar habitats in Eastern parts of Europe outside Russia. On the basis of existing information, this species is undoubtedly acutely threatened within the EU and potential breeding sites should be mapped, in Southern Finland aswell as in other Baltic regions. The distribution of Hylochares spp. across the Holarctic in combination with their habitat choice suggests they are one of the few eucnemids adapted into a life in Northern latitudes.

Dirrhagofarsus ernae Otto, Muona, and McClarin, new species (Figures 1���5) Diagnosis. Within North American Dirhagini, characterized by lateral carinae on frons combined with simple elytral apex. Description. Adult. Holotype male. Length 5 mm; greatest width 1.5 mm. Color black with anterior pronotal margin red-brown. Form subcylindrical, moderately elongate and tapering towards apex (Fig. 1), dorsum clothed with vestiture of short, yellow decumbent hairs. Head subspherical with convex, smooth, shiny and punctate frons. Lateral frontal keels sharp, short, slightly bent and covered with micro-combs, giving a silvery appearance. Vertex with undulating, complete belt of micro-combs. Mandibles stout, bidentate and densely punctate. Antennae redbrown, evenly colored and longer than half the length of the body. Antennomere 3 about 1.4 times as long as 4, antennomeres 4���10 subtriangular and elongated. Antennomere 11 at least 2 �� times longer than wide. Pronotum slightly transverse and subquadrangular. Anterior portion of pronotum slightly convex. Surface shiny, with moderately dense punctures; small and slightly spaced, sides parallel. Disc of pronotum with a slight depression on both sides. Anterior lateral pronotal carinae reach about �� the length of pronotum, while the posterior lateral pronotal carinae (Fig. 2) reach more than �� the length of pronotum. Notosternal antennal grooves shiny and smooth. Inner ridge of antennal grooves obliterated on apical half. Scutellum densely punctate, subtriangular and narrowly rounded at end. Elytra have strongly punctate striae and interspaces. Punctures are partly confused, especially basally. Interspaces are convex at the humeral region and largely flat elsewhere on elytra. Legs redbrown in color. Tibiae rounded in cross section. Lateral surfaces of meso- and meta-tibae with single spines. Claws simple. Tarsomere 4 excavate emarginate. Tarsomeres 1 to 3 simple, protarsomere 1 without sex-comb. Tarsomere 1 as long as the preceding four combined in meso- and meta-tarsi. Metatarsomere 2 twice as long as third and as long as fifth. Aedeagus similar to that of D. attenuatus (see Burakowski 1991), but secondary lateral lobes shorter, lateral lobes apically sharper and median lobe slightly shorter (Fig. 4). Aedeagus also similar to that of D. modestus (Fig. 5), but secondary lateral lobes reach nearly the apex of the aedeagus in D. modestus and the apical part of the organ is proportionately smaller and wider, the basal part elongated and narrow. The flagellum in D. ernae is slender and with small hooks evenly distributed towards the apex, whereas in D. modestus it is medially wide and the hooks are more numerous and absent from the apical region. We have examined 68 adult paratypes. Nine larvae and three pupae were also studied. Females are on average slightly larger than males. Their antennae are stouter, not quite reaching half the length of the body, antennomere 11 being about two times longer than wide (Fig. 3). The pronotum is mostly densely punctate to rugose, slightly dullish. The posterior lateral pronotal carina is shorter than in the males, reaching about 2 / 3 the length of the pronotum. The anterior lateral carina is slightly more acutely angled in the females than in the males. This pronotal dimorphism seems to be true for D. modestus and D. attenuatus as well. The length and height of median carina near the posterior end of the pronotum is variable, many specimens exhibiting a small, shallow to nearly indistinct median carina. Color intensity along the anterior margin of pronotum is slightly variable, but only a few males show a uniform black pronotum, typical of D. modestus. Holotype: Ohio, Hamilton Co., male, ���OH: Hamilton Co., Cincinnati, Mount Airy Forest, May 17, 1998, Eric Eaton coll���. Paratypes: Alabama, Dale Co.: 1 example, ���Ala. Dale Co., Ft. Rucker Mil. Res., sugar bait, 1���7 May 1999, R. Turnbow���; 1 example, ���Ala. Dale Co., Ft. Rucker Mil. Res., 5���12 may 2009, R. Turnbow, manuka oil baited Lindgren funnel trap���. Missouri, DeKalb Co.: 1 example, ���MO: DeKalb Co., 13.4 mi. Cameron, 25 May 2007, R. Turnbow���; Stone Co.: 1 example, ���MO: Stone Co., Ashe Juniper Natural Area, 4.0 mi. N Hwy 86 on JJ, T 22 N R 22 W sec. 20, T.C. MacRae, 00- 90 b: emerged. 16���23.v. 2001, ex. 1���3 " diameter dead branch Acer saccharum L., coll. 17.ix. 2000 ���. New Hampshire, Hillsborough Co.: 26 examples (14 adults, 9 larvae and 3 pupae), ���NH: Hillsborough Co., Nashua, 10 Jan, 2007, Jim McClarin, Reared from larvae in white rotten aspen���. Ohio, Hamilton Co.: 6 examples, ���OH: Hamilton Co., Cincinnati, Mount Airy Forest, May 17, 1998, Eric Eaton Coll.���; 1 example, ���OH: Hamilton Co., Cincinnati, Price Hill, June 24, 1998, Eric Eaton coll.���; Preble Co.: 2 examples, ���OH: Preble Co., Camden, 6 June 2001, M. Ulyschen���; Butler Co.: 1 ex., ���OH: Butler Co., Oxford, 6 July 2001, M. Ulyschen���. Pennsylvania, Westmoreland Co.: 2 examples, ��� USA, PA, Westmoreland Co., 10 KM S. Ligonier, Powdermill Nat. Res., 29 VI 1996; dead trees and fungus at night; D.A. Pollock���. Virginia, Montgomery Co.: 3 examples, ���VA: Montgomery Co., ca 3 km SE Ellett, off Rt. 641, Slaughterpen Hollow, DenCrk, 10 June 2001, J.C. Ludwig UV���; Floyd Co.: 1 example, ���VA: Floyd Co., Buffalo Mtn., 3500 feet, 3500���3900 ', 20 June 2001, S.M. Roble UV���; Bath Co.: 1 example, ���VA: Bath Co., Warm Springs Mtn., Jct. Ridgecut Rd & Rd to Trappers Lodge, 14 June 1999, old growth nr. clear cut, J.C. Ludwig UV���; 1 example, ���VA: City of Richmond, 9404 Derbyshire Road, Apr���Aug 1995, W.H. Mitchell���. Wisconsin, Dane Co.: 1 example, ���NG 14113, United States of America, Wisconsin, Dane County, Madison, Eagle Heights, (300 m), 43 �� 05' N 89 �� 26 ' W, 12.VII. 2003, Gompel N. Leg.���; La Crosse Co.: 1 example., ���WI: La Crosse Co., West Salem, Site 8, N 44 �� 3.589 ' W 91 �� 4.430 ', root sever, Lindgren trap, 29 June 2004, Robert Murphy���; 1 ex., ���WI: La Crosse Co., West Salem, Site 3 B, N 44 �� 4.082 ' W 91 �� 4.575 ', root sever, flight intercept trap, 26 July��� 9 August 2004, Robert Murphy���; 1 example., ���WI: La Crosse Co., West Salem, Site 8, N 44 �� 3.589 ' W 91 �� 4.430 ', root sever, flight intercept trap, 26 July��� 9 August 2004, Robert Murphy���; 1 example., ���WI: La Crosse Co., West Salem, Site 2, N 44 �� 3.411 ' W 91 �� 3.430 ', pkt control, flight intercept trap���inside, 23 June 2005, Robert Murphy���; 2 examples., ���WI: La Crosse Co., West Salem, Site 1, N 44 �� 3.778 ' W 91 �� 4.332 ', pkt control, flight intercept trap���inside, 7 July 2005, Robert Murphy���; 2 examples., ���WI: La Crosse Co., West Salem, Site 5, N 44 �� 3.963 ' W 91 �� 4.521 ', root sever, flight intercept trap���outside, 7 July 2005, Robert Murphy���; 1 example., ���WI: La Crosse Co., West Salem, Site 2, N 44.06296 �� W 91.07220 ��, pkt control, flight intercept trap���inside, 21 July 2005, Robert Murphy���; 1 example., ���WI: La Crosse Co., West Salem, Site 5, N 44.06605 �� W 91.07220 ��, root sever, flight intercept trap���outside, 21 July 2005, Robert Murphy���; Milwaukee Co.: 1 example., ���NG 14074, United States of America, Wisconsin, Milwaukee County, South Milwaukee, Grant Park, Seven Bridges, 42 �� 55 ' N 87 �� 50 ' W, Woods, 2.VII. 2003, Gompel N. Leg.���; Sauk Co.: 1 example., ���WI: Sauk Co., Gatshall property, N 43 �� 4.765 ' W 89 �� 34.576 ', asystematic control, flight intercept trap���inside, 16 June 2005, Robert Murphy���; 1 example., ���WI: Sauk Co., Bakken's Pond, Site 2, N 43 �� 10.830 ' W 90 �� 9.205 ', pkt. control, flight intercept trap���out, 16 June 2005, Robert Murphy���; Walworth Co.: 1 example., ���WI: Walworth Co., Kettle Moraine St. Forest, N 42 �� 49.551 ' W 88 �� 36.583 ', asystematic control, flight intercept trap���inside, 15 June 2005, Robert Murphy���; 2 examples., ���WI: Walworth Co., Kettle Moraine St. Forest, N 42 �� 49.610 ' W 88 �� 36.471 ', root sever, flight intercept trap���outside, 28 June 2005, Robert Murphy���; 1 example., ���WI: Walworth Co., Kettle Moraine St. Forest, N 42 �� 49.551 ' W 88 �� 36.583 ', asystematic control, flight intercept trap���inside, 28 June 2005, Robert Murphy���; Waukesha Co.: 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 51.574 ' W 88 �� 29.781 ', pkt control, Lindgren Trap, 9 June 2004, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.616 ' W 88 �� 27.891 ', asystematic control, flight intercept trap���inside, 15 June 2005, Robert Murphy���; 2 examples., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.574 ' W 88 �� 29.781 ', pkt control, flight intercept trap���out, 15 June 2005, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 51.574 ' W 88 �� 29.781 ', pkt control, flight intercept trap���inside, 15 June 2005, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.840 ' W 88 �� 28.088 ', root sever, flight intercept trap���out, 15 June 2005, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 49.551 ' W 88 �� 36.583 ', asystematic control, flight intercept trap���outside, 28 June 2005, Robert Murphy���; 4 examples., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.616 ' W 88 �� 27.891 ', asystematic control, flight intercept trap���inside, 29 June 2005, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.840 ' W 88 �� 28.088 ', root sever, flight intercept trap���inside, 29 June 2005, Robert Murphy���; 1 example., ���WI: Waukesha Co., Kettle Moraine St. Forest, N 42 �� 54.840 ' W 88 �� 28.088 ', root sever, flight intercept trap���outside, 29 June 2005, Robert Murphy���; Waupaca Co.: 1 example., ���WI: Waupaca Co., J. Baker property, Site 3, N 44 �� 18.805 ' W 89 �� 6.238 ', asystematic control, flight intercept trap���inside, 22 June 2005, Robert Murphy���. The holotype male and one female paratype are deposited at the US National Museum in Washington DC. Further paratypes are deposited in the American Museum of Natural History, the UW Entomology Museum (UW- Madison), the Insect Research Collection (South Dakota State University), the Field Museum of Natural History in Chicago, the Royal Ontario Museum in Quebec, the Finnish Museum of Natural History, Bohart Museum of Entomology (UC-Davis), Florida State Collection of Arthropods (Gainesville) and Virginia Museum of Natural History as well as the collections of the authors, Eric Eaton, Ed Riley, Ted MacRae and Nicolas Gompel. Etymology. Dirrhagofarsus ernae is named in honor of the senior author���s grandmother, Erna (Karstedt) James who passed away in 1999. Larva. (Figs. 6 ���11) Mature larvae (Fig. 6). Length 14���16 mm, width 1.0��� 1.2 mm. Body subcylindrical, near parallel-sided, pro- and meso-thorax wider than remaining segments, cream-yellow in color with head, prothoracic scleromes and posterior end of ninth abdominal segment dark brown. Head dorsoventrally flattened, broadly subtriangular, prognathus and exerted from prothorax. The sclcerotized cephalic discs form a compact structure with lateral toothlike projections (Figs. 7, 8). Dorsal cephalic disc subtrapezoidal with longitudinal furrows on either side of midline. Longitudinal furrows converge anteriad, forming a small trapezoidal ridge with basal median carina. Ventral cephalic disc pentagonally shaped and with a small single raised carina. Anterior portion of head heavily sclerotized near mandibles. Mandibles heavily sclerotized, oval, longer than wide, deeply bifid, fused to head, resting in mesal acumination of head capsule. Labium weakly chitinized, retractile, with three segments. Labial palps slightly longer with three segments. Visible ligula, mala, lacinia and galea absent. Hypostomal rods also absent. Antennae minute, arising between first and second lateral projections. Basal segment about as long as wide, second segment of equal length, but narrow, with three apical setae. Sensory papillae well developed, about as long as second antennal segment. Prothorax slightly longer than metathorax and abdominal segments subequal to mesothoracic segment, metathorax being shortest of all. Dorsally, near anterior end, a pair of horizonal lineal scleromes extend from lateral sides to mesal ecdysial line, diverge and run length of pronotum. A small circular sclerome present near anterior end of prothorax, laterad of each narrowed sclerome. Ninth abdominal sternite (Fig. 10) heavily sclerotized with prominent semi-circular ring of asperities above anal region. Smaller asperities form oblong, oval ring around the anus and extend to each end of larger spicules. Ninth abdominal tergite (Fig. 9) with well-developed microtrichial patch. Posterior margin of ninth segment sparsely punctate with shallow furrows, strongly sclerotized. Spiracles are annular biforous. The arrangement of the microtrichial patches and areoles on dorsum are shown in Fig. 11. The detailed description of the larva of the Palaearctic Dirrhagofarsus attenuatus (M��klin) (Burakowski, 1989) allows a useful comparison of the larval types.Although superficially similar, the larvae differ in many details. The strongly sclerotized apex of the 9 th segment is pointed and nearly triangular in D. ernae, less sclerotized and abruptly cut off in D. attenuatus. The head capsule of D. attenuatus is longitudinally grooved, especially strongly dorsally, whereas in D. ernae it is clearly smoother. The ventral supporting rods on the prothorax have long laterally diverging cranial tips in D. ernae, whereas in D. attenuatus the cranial tips of these rods are very short. Additional information. The first larva of this species was collected independently of senior authors��� adult finds on January 7, 2007 by J. McClarin in Nashua, New Hampshire, USA. It was found in a fairly dry, long-dead, 1.5 m trunk section of Populus that had broken away from the tree several meters off the ground and become caught in a tangle of Celastrus (bittersweet vines), suspended about 1.5 m off the ground. Over 20 more larvae from 6 mm to 18.5 mm long were collected January 10 th in a return visit to the same trunk section, over half of which was softened by white-rot fungus. The larvae were aligned with the wood grain and many were doubled back in their burrows with head in close proximity to its terminal tergite. In attempt to rear an adult for identification, McClarin placed a majority of freed larvae in a sealed plastic container together with small scraps from the Populus trunk section. Some larger wood chunks were also included in hope they contained additional larvae. While most of the freed larvae did not establish new burrows in the soft wood, many had pupated by Feb. 3 and the first adult eclosed on February 9, followed by many more in later days. Because the species proved unrecognizable from photos, first larvae then adults and pupae were preserved and sent to Muona for identification and subsequent DNA sequencing. Muona had already heard from Otto about his discovery of an apparent undescribed species, and he and Otto eventually determined through their independent finds that they were of the same species. Many photos of larvae, pupae and adults of this species may be seen in the Dirrhagofarsus section of bugguide.net: http://bugguide.net/node/view/ 114573 /bgimage. FIGURE 9. Dirrhagofarsus ernae n. sp., larva, 9 th abdominal segment, dorsal (upper) view. FIGURE 10. Dirrhagofarsus ernae n. sp., larva,, 9 th abdominal segment, ventral (lower) view. FIGURE 11. Dirrhagofarsus ernae n. sp., larva, position of microtrichial patches and areoles on dorsum., Published as part of Otto, Robert L., Muona, Jyrki & Mcclarin, Jim, 2014, Description of Dirrhagofarsus ernae n. sp. with a key to the known Dirrhagofarsus species (Coleoptera: Eucnemidae), pp. 179-184 in Zootaxa 3878 (2) on pages 180-183, DOI: 10.11646/zootaxa.3878.2.4, http://zenodo.org/record/231051, {"references":["Burakowski, B. (1991) Klucze do Oznaczania Owadow Polski. Czesc XIX. Chrzaszcze - Coleoptera. Zeszyt 35 - 37. Cerophytidae, Eucnemidae, Throscidae, Lissomidae. Polskie Towarzystwo Entomologiczne, Warsaw, 91 pp.","Burakowski, B. (1989) Hypermetamorphosis of Rhacopus attenuatus (Maklin) (Coleoptera: Eucnemidae). Annales Zoologici, 42,165 - 180."]}

Otto, Robert L., Muona, Jyrki, Mcclarin, Jim (2014): Description of Dirrhagofarsus ernae n. sp. with a key to the known Dirrhagofarsus species (Coleoptera: Eucnemidae). Zootaxa 3878 (2): 179-184, DOI: http://dx.doi.org/10.11646/zootaxa.3878.2.4

Dirrhagofarsus Fleutiaux, 1935 Type-species: Microrhagus lewisi Fleutiaux. (= Attenuorhagus Olexa, 1975. Type-species Hypocaelus attenuatus M��klin) Diagnosis. (Tribe Dirhagini) Head with transverse belt of microcombs on vertex, lateral keels on frons, deep sensory pits on front margin of large eyes, interrupted lateral pronotal keels, well-developed notosternal antennal grooves with cranially obliterated median ridge, simple male protarsomere 1, metacoxal plates abruptly narrowing laterad, acutely produced apex of last ventrite, male aedeagus with well-developed secondary lateral lobes, acute, apically narrowing lateral lobes, bridged, bifid, narrow median lobe apex and no ventral plate. The aedeagus of D. lewisi differs from those of the other species by the abruptly constricted apical region; secondary lateral lobes are of the same length as those of D. modestus. Terminology follows Muona (1993) and Muona & Ter��v��inen (2009). The trans-Palaearctic and very rare D. attenuatus (M��klin) is a complex of species. We have included the true D. attenuatus in the key on the basis of the syntypes JM studied., Published as part of Otto, Robert L., Muona, Jyrki & Mcclarin, Jim, 2014, Description of Dirrhagofarsus ernae n. sp. with a key to the known Dirrhagofarsus species (Coleoptera: Eucnemidae), pp. 179-184 in Zootaxa 3878 (2) on pages 179-180, DOI: 10.11646/zootaxa.3878.2.4, http://zenodo.org/record/231051, {"references":["Muona, J. (1993) Review of the phylogeny, classification and biology of the family Eucnemidae (Coleoptera). Entomologica Scandinavica Supplement, 44, 1 - 133."]}

Dirrhagus lindbergi Palm, 1958 is regarded to be a junior subjective synonym of Microrhagus lepidus Rosenhauer 1847. The type series of D. lindbergi consisted of two species, the holotype male being conspecific with Microrhagus lepidus, the two paratype females with Microrhagus pygmaeus (Fabricius).

The adult and the larva of Dirrhagofarsus ernae n. sp. are described from the Eastern United States. The genus Dirrhagofarsus is diagnosed and a key to all known species of the genus is provided.

Eighty-nine eucnemid genera occur in the region from South-east Asia to the south-west Pacific. The phylogenies of 84 of these were used together with the present-day distributions of the species to analyse the biogeographical history of the area. Fifty-seven genera shared a pattern coinciding with the traditional model of Laurasia–Gondwana break-up. Six genera showed a pattern contradicting the model. The remaining 21 genera neither supported nor refuted the model. Twenty-five genera were observed to include an Indomalesian clade younger than the South America–Australia connection. This biogeographical unit consisted of present-day South-east Asia and the Sunda islands, but did not include the Philippine Islands and Sulawesi. In addition to this Indomalesian clade, three separate clades involving northern Australia or New Guinea were observed, New Guinea–Australia, New Guinea–Philippines–Sulawesi and New Guinea–Fiji. The possible presence of four separate areas in the general region of New Guinea–north Australia as the result of the Cretaceous geological events is suggested. Three of these, in the area of present-day New Guinea, originally sharing sister-groups with the north-eastern Australian isolate, are regarded as the sources of the New Guinea–Indomalesia, New Guinea–Philippines and New Guinea–Fiji faunas after northward drifting of the Australian continent. During the Oligocene–Miocene these source areas were flooded and their original fauna lost. When the present-day New Guinea emerged, it was invaded from the north-eastern Australian region. This invasion created new New Guinea–Australia connections and brought in the sister-groups of the old New Guinea source areas as well. The eucnemids of Vanuatu, Samoa and Tonga are regarded as having originated in connection with dispersal from Fiji. The New Zealand fauna has strong, old connections with that of south-eastern Australia, but other complex connections are indicated. The Eocene Baltic Amber fauna agrees well with the results obtained from extant species. The species belonging to five fossil genera belong to Gondwanan groups that seem to have invaded the Holarctic via Central America. Four other fossil genera showing discordant patterns belong to the group of six genera exhibiting these aberrant patterns even today. The eucnemid fauna of the region is of Gondwanic origin. Only six Laurasian genera have invaded the area, all of them apparently quite recently.

An alphabetical genus-group catalogue of the family Eucnemidae is given with type species citations. The catalogue is up-dated to 1984. Type designations are made for three names, Compsocnemis Bonvouloir, Feaía Fleutíaux and Sassanus Fleutiaux. The author of the genus Perothops is shown to be Laporte, 1838, that of the genus Oisocerus Bonvouloir in Murray, 1868. The descriptions of Rhigmaphorus Guérin-Méneville, 1843 and Sphaerocephalus brasiliensis, Germar 1839 are regarded to be valid. Haywardinia Cobos, 1964 and Suareziella Cobos, 1964 are shown to be junior homonyms. Willinkia Cobos is shown to be a nomen nudum. The correct usage of some misapplied names is emphasized: Farsus Jacquelin du Val, 1863 (=Hylochares auctt. nec Latreille, 1834), Galbites Fleutiaux, 1918 (=Pterotarsus auctt. nec Guérin-Méneville, 1831). Hylis des Gozis, 1886 (=Hypocaelus auctt. nec Dejean, 1833), Microrhagus Dejean, 1833 (=Dirhagus Latreille, 1834) and Xylophilus Mannerheim, 1823 (=Xyloecus Dejean, 1833, =Xylobius Latreille, 1834).

The taxonomy of the African Arrhipis species is discussed. A. senegalensis (Laporte de Castelnau) is regarded a valid species. Lectotype is designated for A. vassei Fleutiaux. A key including the eight known African species is given as well as data on their distribution.

Phlegon degallieri sp.n. is described from French Guiana. A key to the species of the genus Phlegon is given as well as notes on their distribution.

The larvae of Anischia Fleutiaux and Perothops Laporte are described. Cladistic analyses based on adult and larval morphology, as well as CO1 sequence data, place both genera in the Eucnemidae clade within the Elateroidea (sensu stricto). The subfamily Anischiinae Fleutiaux, 1936 is placed in the family Eucnemidae in a clade containing the more derived eucnemid subfamilies (Melasinae, Eucneminae and Macraulacinae), while Perothops and Phyllocerus Lepeletier & Serville represent subfamilies basal to the remaining eucnemid taxa. The genus Afranischia Basilewsky, 1955 is synonymized with Anischia Fleutiaux, 1896, and Anischia boliviana Fleutiaux is selected as the type species of the latter. Three new species are described: Anischia bicolor (New Caledonia), Anischia kuscheli (New Caledonia) and Anischia monteithi (NE Australia), and Anischia stupenda Fleutiaux, 1897 is recorded from Australia. Anischia crassicornis Champion, 1897 is synonymized with Anischia mexicana Fleutiaux, 1896. One new combination is made, Anischia ruandana (Basilewsky, 1955).

The genus Euyostus Bonvouloir is revised. Two new species are described: E. kuchinensis n.sp. (Myanmar) and E. moorei n. sp. (“Borneo”). Lectotypes are designated for two species: E. reichei Bonvouloir, 1871 (Singapore) and E. bonvouloiri Fleutiaux, 1912 (New Guinea). The habitus and the male genitalia of all the species are illustrated and a key to identify the species is given.

Eight new genera and twenty new species of Eucnemidae are described from Cretaceous Myanmar amber: Cupressicharis n. gen. (type-species Cupressicharis elongatus n. sp.), Cylus n. gen. (type species Cylus carinifer n. sp.), Falsothambus n. gen. (type species Falsothambus burmensis n. sp.). Fiegelia n. gen. (type species Fiegelia antennata n. sp.), Paleoeucnemis n. gen. (type species Paleoeucnemis minutus n. sp.), Protomicrorhagus n. gen. (type species Protomicrorhagus antennatus n. sp.), Protovitellius n. gen. (type species Protovitellius deceptus n. sp,) and Pseudomyall n. gen. (type species Pseudomyall elongatulus n. sp). Eleven further species are described: Coenomana brevicornis n. sp., Euryptychus acutangulus n. sp., Euryptychus burmensis n. sp, Euryptychus elegantulus n. sp., Euryptychus mysticus n. sp, Epiphanis burmensis n. sp., Falsothambus gracilicornis n. sp., Fiegelia tarsalis n. sp., Jenibuntor pusillus n. sp., Myall burmensis n. sp., Protomicrorhagus brevis n. sp, Sieglindea antiqua n. sp. Coenomana clavata Otto is transferred from tribe Jenibuntorini to tribe Dirhagini. Muonabuntor grandinotalis Li , Tihelka & Cai is transferred from tribe Jenibuntorini to tribe Euryptychini.