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Background: Recently, common genetic risk factors for intracranial aneurysm (IA) and aneurysmal subarachnoid hemorrhage (ASAH) were found to explain a large amount of disease heritability and therefore have potential to be used for genetic risk prediction. We constructed a genetic risk score to (1) predict ASAH incidence and IA presence (combined set of unruptured IA and ASAH) and (2) assess its association with patient characteristics. Methods: A genetic risk score incorporating genetic association data for IA and 17 traits related to IA (so-called metaGRS) was created using 1161 IA cases and 407 392 controls from the UK Biobank population study. The metaGRS was validated in combination with risk factors blood pressure, sex, and smoking in 828 IA cases and 68 568 controls from the Nordic HUNT population study. Furthermore, we assessed association between the metaGRS and patient characteristics in a cohort of 5560 IA patients. Results: Per SD increase of metaGRS, the hazard ratio for ASAH incidence was 1.34 (95% CI, 1.20–1.51) and the odds ratio for IA presence 1.09 (95% CI, 1.01–1.18). Upon including the metaGRS on top of clinical risk factors, the concordance index to predict ASAH hazard increased from 0.63 (95% CI, 0.59–0.67) to 0.65 (95% CI, 0.62–0.69), while prediction of IA presence did not improve. The metaGRS was statistically significantly associated with age at ASAH (β=−4.82×10 −3 per year [95% CI, −6.49×10 −3 to −3.14×10 −3 ]; P =1.82×10 −8 ), and location of IA at the internal carotid artery (odds ratio=0.92 [95% CI, 0.86–0.98]; P =0.0041). Conclusions: The metaGRS was predictive of ASAH incidence, although with limited added value over clinical risk factors. The metaGRS was not predictive of IA presence. Therefore, we do not recommend using this metaGRS in daily clinical care. Genetic risk does partly explain the clinical heterogeneity of IA warranting prioritization of clinical heterogeneity in future genetic prediction studies of IA and ASAH.

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

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.)"]}

Data de publicació electrònica: 04-04-2022 Pathogenic variants in A Disintegrin And Metalloproteinase (ADAM) 22, the postsynaptic cell membrane receptor for the glycoprotein leucine-rich repeat glioma-inactivated protein 1 (LGI1), have been recently associated with recessive developmental and epileptic encephalopathy. However, so far, only two affected individuals have been described and many features of this disorder are unknown. We refine the phenotype and report 19 additional individuals harboring compound heterozygous or homozygous inactivating ADAM22 variants, of whom 18 had clinical data available. Additionally, we provide follow-up data from two previously reported cases. All affected individuals exhibited infantile-onset, treatment-resistant epilepsy. Additional clinical features included moderate to profound global developmental delay/intellectual disability (20/20), hypotonia (12/20), delayed motor development (19/20). Brain MRI findings included cerebral atrophy (13/20), supported by post-mortem histological examination in patient-derived brain tissue, cerebellar vermis atrophy (5/20), and callosal hypoplasia (4/20). Functional studies in transfected cell lines confirmed the deleteriousness of all identified variants and indicated at least three distinct pathological mechanisms: defective cell membrane expression (1), impaired LGI1-binding (2), and/or impaired interaction with the postsynaptic density protein PSD-95 (3). We reveal novel clinical and molecular hallmarks of ADAM22 deficiency and provide knowledge that might inform clinical management and early diagnostics. Funding: most families were collected as part of the SYNaPS Study Group collaboration funded by the Wellcome Trust and strategic award (Synaptopathies) funding (WT093205 MA and WT104033AIA). This study was also supported by JSPS/MEXT KAKENHI (Grants 19H03331, 19K22439 and 21K19390 to Y.F., Grant 19K16269 to Y.M., and Grants 20H00459 and 20H04915 to M.F.) and Japan Agency for Medical Research and Development (21wm0525022h0001 to Y.F.); intramural funding (fortüne) from the University of Tübingen (Grant 2545-1-0) and the Ministry of Science, Research and Art Baden-Württemberg to B.V. P.S. contributed to this work within the framework of the DINOGMI Department of Excellence of MIUR 2018-2022 (legge 232 del 2016). T.B.H. was supported by the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation) – 418081722, 433158657. I.S.P., V.Y.V. are supported by the Government Assignment of the Russian Ministry of Health (#121061500066-2). HL receives support from the Canadian Institutes of Health Research (Foundation Grant FDN-167281), the Canadian Institutes of Health Research and Muscular Dystrophy Canada (Network Catalyst Grant for NMD4C), the Canada Foundation for Innovation (CFI-JELF 38412), and the Canada Research Chairs program (Canada Research Chair in Neuromuscular Genomics and Health, 950-232279). S.H. is funded by TUBITAK (Turkish Scientific and Technological Research Council) Project number 216S771. R.H. is a Wellcome Trust Investigator (109915/Z/15/Z), who received support from the Medical Research Council (UK) (MR/N025431/1 and MR/V009346/1), the European Research Council (309548), the Newton Fund (UK/Turkey, MR/N027302/1), the Addenbrookes Charitable Trust (G100142), the Evelyn Trust, the Stoneygate Trust, the Lily Foundation and an MRC strategic award to establish an International Centre for Genomic Medicine in Neuromuscular Diseases (ICGNMD) MR/S005021/1. This research was supported by the NIHR Cambridge Biomedical Research Centre (BRC-1215-20014)