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Dyscolus silvestris Moret sp. nov. urn:lsid:zoobank.org:act: 8DA5B9B7-F332-4D75-AC2D-CEBBD34ADA3C Figs 52���53 Etymology Latin adjective meaning ���forest related���. Type material Holotype ECUADOR ��� ♂; Napo Province, Termas de Papallacta; 3300 m a.s.l.; 1 Jul. 2001; P. Moret leg.; ���sousbois���; QCAZ. Paratypes (5 ♂♂, 10 ♀♀) ECUADOR ��� Napo Province ��� 1 ♂, 1 ♀; same collection data as for holotype; CPM ��� 1 ♂, 1 ♀; Papallacta, Termas Jamanco, Waypoint 41; 0��22���24.6��� S, 78��10���5.5��� W; 3410 m a.s.l.; 25 Oct. 2015; P. Moret leg.; MNHN ��� 1 ♀; same collection data as for preceding; QCAZ ��� 1 ♂, 3 ♀♀; same collection data as for preceding; CPM ��� 1 ♂; same collection data as for preceding; COI voucher PM041-02, BOLD sequence SUM103-18; CPM ��� 1 ♀; same collection data as for preceding; COI voucher in ethanol PM041-06, BOLD sequence SUM104-18; CPM ��� 2 ♀♀; Lago Papallacta; 3370 m a.s.l.; 9 Apr. 1986; A. Vigna; CPM ��� 1 ♀; Papallacta; 2700 m a.s.l.; 6���8 Apr. 1979; H. Frania leg.; UASM ��� 1 ♂; Lago Papallacta; 0��20���29��� S, 78��10���23��� W; 3400 m a.s.l.; 6 Nov. 1999; R. Anderson leg.; CMNC. Diagnostic description Habitus: Fig. 52. Wingless. Body length: 9.1���10.0 mm. Body colour variable, predominantly dark brown with reddish brown areas in some specimens; legs, antennae and mouthparts reddish brown. Elytral microsculpture isodiametric. Head broad and convex, neck slightly constricted, eyes moderately bulging. Pronotum subcordiform, the base much narrower than the apex; sides arcuate anterad, markedly sinuate posterad; hind angles obtuse, sharp; two pairs of lateral setae. Elytra subovate, moderately convex; striae entire, well impressed; intervals flat. Third elytral interval with 3���4 discal setae. Legs: protarsi of the male much broader than those of the female. Fifth metatarsomere asetose ventrally. Last visible abdominal ventrite with one pair (♂) or two pairs (♀) of setae along its apical margin. Male genitalia: Fig. 53. Median lobe arcuate, shortly and bluntly triangular at apex, endophallus unarmed. Female genitalia: unstudied. Comparisons Dyscolus silvestris Moret sp. nov. is similar to the shorter and broader individuals of Dyscolus denigratus. The main differences are: head slightly bigger and broader; sides of the pronotum sinuate basally, hind angles sharply obtuse (completely rounded in D. denigratus); femora of the male not swollen as in D. denigratus. The apex of the median lobe of the male genitalia has a shorter apex, and the endophallus is entirely unarmed (with a heavily sclerotized area in D. denigratus). Habitat Upper montane forest on the Amazonian slope of the Andes, from 2700 to 3400 m a.s.l. This species lives in the leaf litter. Geographic distribution Only known from the type locality around Papallacta in Northern Ecuador, probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 45-47, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus arborarius Moret sp. nov. urn:lsid:zoobank.org:act: 6FD45E2B-C31E-4486-8285-F537FCBF4FB6 Figs 25���26 Etymology Latin adjective meaning ���related to trees���. Type material Holotype (only known specimen) ECUADOR ��� ♂; Loja Province, Parque Nacional Yacuri, Waypoint 170; 4.781387�� S, 79.386938�� W; 2870 m a.s.l.; 4 Aug. 2016; P. Moret and S. Aguirre leg.; COI voucher PM170-02, BOLD sequence SUM191-18; MNHN. Diagnostic description Habitus: Fig. 25. Wingless. Body length: 10.8 mm. Body dark brown with the sides of the pronotum and the elytral sutura reddish brown; femora dark brown; rest of the legs, antennae and mouthparts reddish brown. Elytral microsculpture shallowly impressed, transverse. Head large and convex; eyes prominent, very convex; basal constriction strong; mandibles relatively short. Antennae thin, very elongate. Pronotum subcordiform, sides moderately arcuate anterad, slightly sinuate posterad; hind angles obtuse; two pairs of lateral setae. Elytra elongate-ovate with effaced humeri, subapical sinuation obsolete; striae narrow and shallowly impressed; intervals flat; third elytral interval with two or three discal setae. Last visible abdominal ventrite of the male with two pairs of apical setae. Legs thin and elongate. Fourth metatarsomere oval-shaped, with one pair of dorsolateral subapical setae and long apical lobes, the external lobe nearly two times longer than the inner lobe. Male genitalia: Fig. 26. Median lobe weakly arcuate, apex short, acute, endophallus with one subapical sclerotized structure. Female genitalia: unknown. Comparisons According to the COI gene, D. arborarius Moret sp. nov. belongs to the marini clade, close to D. ruizi Moret sp. nov. It can be easily distinguished from the other species of this clade by its slender body, large head with very prominent eyes, and effaced humeri. Habitat Upper montane forest on the Eastern slope of the Andes, at around 2870 m a.s.l. Collected by pyrethrin insecticide fogging on mossy branches and epiphytes, two metres above ground. Geographic distribution Only known from the type locality in Southern Ecuador, in Parque Nacional Yacuri. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 27-29, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus famelicus Moret sp. nov. urn:lsid:zoobank.org:act: FC15AE30-F921-48BF-BFDE-DD80F0CF7351 Figs 27���28 Etymology Latin adjective meaning ���scrawny, gaunt���, in allusion to the narrow and slender body of this species. Type material Holotype ECUADOR ��� ♂; Napo Province, East of Papallacta���Guango Lodge, Waypoint 40; 0��22���42.6��� S, 78��4���26.6��� W; 2708 m a.s.l.; 24 Oct. 2015; P. Moret leg.; QCAZ. Paratypes (2 ♂♂, 2 ♀♀) ECUADOR ��� 1 ♂; same collection data as for holotype; MNHN ��� 1 ♂, 1 ♀; same collection data as for holotype; CPM ��� 1 ♀; same collection data as for holotype; COI voucher PM040-02, BOLD sequence SUM184-18; CPM. Diagnostic description Habitus: Fig. 27. Wingless. Body length: 9.6���10.5 mm. Elytra brownish, disc of the pronotum brownish black, margins of the pronotum brownish as the elytra, head piceous black; femora brownish, tibiae, tarsi, mouthparts and antennae reddish brown. Elytral microsculpture isodiametric. Head narrow with bulging eyes, frons with two foveae. Pronotum elongate, as wide as long, constricted basally; lateroapical lobes reduced; hind angles laterally prominent, broadly rounded; two pairs of lateral setae. Elytra elongate, ovoid, with effaced humeri and no hint of subapical sinuation; striae shallow, broken into dashes over elytral disc; intervals flat; third interval without discal setae. Last visible abdominal ventrite with one pair (♂) or two pairs (♀) of setae along its apical margin. Legs and antennae elongate. Male genitalia: Fig. 28. Median lobe forming an obtuse angle after basal third (in lateral view), apex short, denticulate and bent downward in lateral view; endophallus without sclerotized structure. Female genitalia: unstudied. Comparisons The slender, elongate body form, the dashed elytral setae and the hooked apex of the aedeagus separate D. famelicus Moret sp. nov. from any other species having an asetose third elytral interval. Our molecular analysis (Fig. 2) places this species close to D. gobbii Moret sp. nov. which lives in the p��ramo and has a very different external morphology, but the bootstrap support for this relationship is weak. Habitat Upper montane forest on the Eastern slope of the Andes, at around 2700 m a.s.l., collected above ground at the beginning of the night, on mossy trunks and on epiphytes. Geographic distribution Only known from the type locality in Northern Ecuador, probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 29-30, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus ravidus Moret sp. nov. urn:lsid:zoobank.org:act: 8449EDAE-130C-468E-A1F3-78EE6FB86649 Figs 30���31 Etymology Latin adjective meaning ���dark���. Type material Holotype ECUADOR ��� ♂; Loja Province, Cajanuma, Refugio Parque Nacional Podocarpus; 4��6���58.4��� S, 79��10���18.6��� W; 2850 m a.s.l.; 17 Mar. 2015; P. Moret and C. Ruiz leg.; QCAZ. Paratypes (7 ♂♂, 5 ♀♀) ECUADOR ��� Loja Province ��� 1 ♂; same collection data as for holotype; CPM ��� 1 ♂; same collection data as for holotype; QCAZ ��� 1 ♂, 1 ♀; Cajanuma, Refugio Parque Nacional Podocarpus, Waypoint 166; 4.11617�� S, 79.17204�� W; 2840 m; 2 Aug. 2016; P. Moret leg.; CPM ��� 1 ♂, 1 ♀; same collection data as for preceding; MNHN ��� 1 ♂; Valladolid, L��mite del Parque Jocotoco y Podocarpus; 6 Jan. 2001; I.G. Tapia leg.; QCAZ ��� 1 ♀; Parque Nacional Podocarpus; 2800 m a.s.l.; 12 Feb. 1994; G. Onore leg.; QCAZ ��� 1 ♀; Mamanuma; 2700 m a.s.l.; 28 Dec. 1994; G. Onore leg.; CISEC ��� 1 ♂; Parque Nacional Podocarpus, Cajanuma, Waypoint 397; 4��7���2.4��� S, 79��10���6.3��� S; 2970 m a.s.l.; 17 Mar. 2015; P. Moret and C. Ruiz leg.; COI voucher PM397-02, BOLD sequence SUM058-18; CPM ��� 1 ♂; same collection data as for preceding; COI voucher PM397-03, BOLD sequence SUM249-18; CPM ��� 1 ♀; km 10 v��a Saraguro���Loja; 2850 m a.s.l.; 21 Jul. 1988; P. Moret leg.; CPM. Diagnostic description Habitus: Fig. 30. Wingless. Body length: 9.5���10.8 mm. Body colour from reddish brown to brunneous; legs, antennae and mouthparts reddish brown. Elytral microsculpture transverse. Head elongate, markedly constricted basally, eyes moderately convex, genae flat. Pronotum subcordiform; sides arcuate anterad, sinuate posterad before the hind angles which are sharp and right angled. Elytra slightly convex; striae entire but narrow and shallowly impressed, intervals flat; third interval without discal setae. Last visible abdominal ventrite with one pair (♂) or two pairs (♀) of setae along its apical margin. Legs: tarsi smooth dorsally, metatarsomeres 1���3 bisulcate, fourth metatarsomere with one pair of dorsolateral subapical setae, apical lobes moderately long, the external lobe> 2 times longer than the inner lobe; fifth metatarsomere asetose ventrally. Male genitalia: Fig. 31. Ventral side of the median lobe bisinuate in lateral view; apical blade elongate; endophallus with a dense squamose area. Female genitalia: unstudied. Comparisons According to the currently available molecular data, Dyscolus ravidus Moret sp. nov. belongs to the same clade as D. crespoae Moret sp. nov. (see above in the description of the latter for the morphological differences between them). However, by its external characters, D. ravidus Moret sp. nov. has a closer resemblance to D. marini Moret sp. nov., which belongs to a different clade. Dyscolus ravidus Moret sp. nov. differs from D. marini Moret sp. nov. by the following characters: elytra brownish in contrast with a black or piceous black pronotum; eyes less convex; hind angles of the pronotum sharper, right angled; elytral striae narrower and more shallowly impressed; third interval without discal setae; elytral microsculpture transverse; male genitalia: apical blade of the median lobe two times as long as in D. marini Moret sp. nov. Habitat Upper montane forest on the Eastern slope of the Andes, between 2700 and 2900 m a.s.l. Active at the beginning of the night (7.30���9 pm) on the ground and in leaf axils of epiphytic bromeliads. Geographic distribution Only known from the Parque Nacional Podocarpus in Southern Ecuador. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 32-33, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus marini Moret sp. nov. urn:lsid:zoobank.org:act: 3E0B11ED-E5E4-438D-9E7B-F7FEC399804B Figs 21, 24 Etymology Noun in the genitive case, dedicated to Diego Mar��n, curator of the insect collection (CISEC) at the Universidad T��cnica Particular de Loja, Ecuador. Type material Holotype ECUADOR ��� ♂; Loja Province, Parque Nacional Podocarpus, Cajanuma, Waypoint 392; 4��6���58.4��� S, 79��10���18.6��� W; 2850 m a.s.l.; 17 Mar. 2015; P. Moret and C. Ruiz leg.; 19.30���21 pm; bromeliad on tree; COI voucher PM392-10, BOLD sequence SUM057-18; QCAZ. Paratypes (5 ♂♂, 3 ♀♀) ECUADOR - Loja Province ��� 1 ♀; same collection data as for holotype; MNHN ��� 1 ♂; Parque Nacional Podocarpus, Cajanuma, Waypoint 397; 4��7���2.4��� S, 79��10���6.3��� W; 2970 m a.s.l.; 17 Mar. 2015; P. Moret and C. Ruiz leg.; COI voucher PM397-01, BOLD sequence SUM248-18; CPM ��� 1 ♀; same collection data as for preceding; QCAZ 252423 ��� 1 ♂; Parque Nacional Podocarpus, Cajanuma, Waypoint 399; 4��7���3.4��� S, 79��9���51.0��� W; 3020 m a.s.l.; 17 Mar. 2015; P. Moret and C. Ruiz leg.; COI voucher PM399- 02, BOLD sequence SUM189-18; CPM ��� 1 ♀; Parque Nacional Podocarpus, Cajanuma; 4.115 S, 79.171 W; 2870 m a.s.l.; 4 Jul. 2013; D. Mar��n and C. Ruiz leg.; COI voucher CR029, BOLD sequence GBNP020-17; CISEC ��� 1 ♂; Parque Nacional Podocarpus, Cajanuma; 4��5���5 S, 79��12��� W; 2750 m a.s.l.; 28���30 Dec. 2001; G. Beltr��n leg.; QCAZ 74257. Diagnostic description Habitus: Fig. 21. Wingless. Body length: 9.9���12.3 mm. Body colour variable, from reddish brown to brunneous; legs, antennae and mouthparts reddish brown. Elytral microsculpture isodiametric. Head elongate, markedly constricted basally, eyes prominent, genae flat. Pronotum subcordiform; sides arcuate anterad, sinuate posterad before the hind angles which are slightly obtuse or right angled; two pairs of lateral setae. Elytra depressed medially; striae entire, well impressed, relatively broad; intervals flat. Third elytral interval with two or three setae (when only two setae, the median one is missing). Last visible abdominal ventrite with one pair (♂) or two pairs (♀) of setae along its apical margin. Legs: tarsi rugose dorsally, metatarsomeres 1���3 bisulcate, fourth metatarsomere with one pair of dorsolateral subapical setae, apical lobes moderately long, the external lobe 1.5 times longer than the inner lobe; fifth metatarsomere asetose ventrally. Male genitalia: Fig. 24. Median lobe arcuate, apex short, endophallus without sclerotized structure. Female genitalia: unstudied. Comparisons According to the currently available molecular data, Dyscolus marini Moret sp. nov. belongs to the same clade as D. ruizi Moret sp. nov., D. arborarius Moret sp. nov., D. moretianus Perrault, 1993, D. sulcipedis Moret sp. nov. and D. rugitarsis Moret sp. nov. However, by its external characters it has a close resemblance to D. ravidus Moret sp. nov., which belongs to a different clade. Dyscolus marini Moret sp. nov. differs from D. ravidus Moret sp. nov. by the following characters: elytra with the same colour as the pronotum; eyes more convex; hind angles of the pronotum obtuse; elytral striae wider; third interval with at least two discal setae; elytral microsculpture isodiametric; male genitalia: apical blade of the median lobe two times shorter than in D. ravidus Moret sp. nov. Habitat Upper montane forest on the Eastern slope of the Andes, 2750���3050 m a.s.l., found at the beginning of the night on the surface of the leaf litter and above ground on mossy trunks or on epiphytes, especially in leaf axils of epiphytic bromeliads. Geographic distribution Only known from the type locality in Southern Ecuador. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 23-24, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus piscator Moret sp. nov. urn:lsid:zoobank.org:act: C207DE54-A6E2-4C10-AF45-EDBE8BAF3D22 Figs 37, 40, 41 (below) Etymology Latin adjective meaning ���fisherman���, in allusion to the feeding habits of this riparian species which preys on small water invertebrates. Type material Holotype ECUADOR ��� ♂; Napo Province, Guaman��, Paso de la Virgen, Waypoint 283; 0��21���0.6��� S, 78��11���52.4��� W; 3890 m a.s.l.; 11 Mar. 2017; P. Moret and M. Gobbi leg.; QCAZ. Paratypes (2 ♂♂, 1 ♀) ECUADOR ��� 1 ♂; same collection data as for holotype; MNHN ��� 1 ♂; same collection data as for holotype; CPM ��� 1 ♀; same collection data as for holotype; COI voucher in ethanol PM283-01, BOLD sequence SUM239-18; CPM. Diagnostic description Habitus: Fig. 37. Wingless. Body length: 9.6���10.9 mm, bigger than D. placitus Moret sp. nov. Ventral surface paler, abdominal ventrites entirely reddish-testaceous. Elytral microsculpture slightly less transverse. Head bigger, genae slightly but distinctly convex in dorsal view (flat in D. placitus Moret sp. nov.); apex of the labrum bisinuate (straight in D. placitus Moret sp. nov.); mandibles much longer, finely acute at apex (length of the right mandible / width of the frons between the anterior setae = 0.95 in D. piscator Moret sp. nov., 1.21 in D. placitus Moret sp. nov.). Third elytral interval with 3���4 setae, fifth interval with 2���4 setae, seventh interval with 0���1 seta; 11���13 umbilicate setae along the lateral margin (17���20 in D. placitus Moret sp. nov.). Male genitalia: Fig. 40. Apex of the median lobe slightly curved, not strongly bent downwards as in D. placitus Moret sp. nov., with a rounded tip, not hooked as in D. placitus Moret sp. nov. Habitat Riparian in the upper montane forest with Polylepis, at around 3900 m a.s.l. Under stones at the edge of a small stream. This biotope is 50 m away from that of D. placitus Moret sp. nov. Geographic distribution Microendemic species, restricted to the Guaman�� area in the Eastern Cordillera., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 37-38, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus globoculus Moret sp. nov. urn:lsid:zoobank.org:act: BDCC6B63-F8CC-4438-8CB3-C7458BCC9715 Figs 13���14 Etymology Combination of two Latin words, ��� globus ��� and ��� oculus ���, in allusion to the globular form of the eyes of this new species. Type material Holotype (only known specimen) ECUADOR ��� ♂ (lacks the left middle leg, the left hind leg and the three last joints of the right antenna); Zamora-Chinchipe Province, Parque Nacional Podocarpus, Bajada Carretera Loja���Zamora; 3.987�� S, 79.118�� W; 2530 m a.s.l.; 5 Sep. 2013; D. Mar��n and C. Ruiz leg.; COI voucher CR064, BOLD sequence GBNP035-17; MNHN. Diagnostic description Habitus: Fig. 13. Wingless. Body length: 12.2 mm. Body shiny black, with a bluish-violaceous metallic lustre on the elytra; femora and tibiae black, tarsi dark brown; palpi reddish brown, antennomeres 1���3 nigropiceous with a darker apex, 4���11 reddish brown. Elytral microsculpture transverse. Head small, very elongate, almost cylindrical, with an abrupt collar constriction; eyes strongly protruding, almost hemispherical; genae long and flat, longer than the eyes; mandibles moderately long and acute. Pronotum almost as wide as the elytra, completely rounded basally, with broadly explanate margins and lacking anterior lobes; two pairs of lateral setae; longitudinal basal impressions narrow. Elytra elongate-oval, convex; humeri effaced; subapical sinuation weak; apex separately rounded. Striae deep and punctate, randomly broken by a few interruptions in the distal part of striae 1���4 and all along striae 5���7. Intervals 1���7 strongly convex, the third interval with two discal setae, one after the middle at ca 0.60 of the elytron length, the other in a broader fovea at 0.85; 17 umbilicate setae along the lateral margin. Last visible abdominal ventrite emarginate, with one pair (♂) of setae along its apical margin. Legs elongate and slender; fourth metatarsomere with one pair of subapical dorsolateral setae (the outer seta has fallen out but the pore is clearly visible), apical lobes asymmetrical, the outer lobe more than two times longer than the inner lobe; fifth tarsomere of all legs setose ventrally. Male genitalia: Fig. 14. Median lobe relatively short, feebly arcuate, apex broadly rounded in dorsal view, endophallus unarmed. Female genitalia: unknown. Comparisons Given the unique combination of the following five characters: elytra with violaceous metallic lustre, cylindrical head with hemispherical eyes, rounded pronotum, bisetose third interval of the elytra and setose onychia, Dyscolus globoculus Moret sp. nov. has no close relatives among Neotropical Platynini. Dyscolus globoculus Moret sp. nov. superficially resembles Javanese members of the genus Colpodes in its strongly protruded eyes and explanate margins of the pronotum (Liebherr 1998: fig. 2), but it lacks several derived characters of Colpodes: head with deep sulci bordering the eyes; effaced neck constriction; elytra with a denticle at the sutural apex; fourth tarsomeres without any subapical dorsolateral seta (Moret 1989: fig. 5; Liebherr 1998: figs 4���6). The protuberant eye is probably a homoplastic character which developed independently in these two lineages. Habitat Upper montane forest on the Eastern slope of the Andes, at around 2500 m a.s.l. Geographic distribution Only known from the type locality in Southern Ecuador, in the Parque Nacional Podocarpus. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 18-20, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682, {"references":["Liebherr J. K. 1998. On Rembus (Colpodes) brunneus MacLeay (Coleoptera: Carabidae, Platynini): redescription and relationships. Journal of Natural History 32: 987 - 1000. https: // doi. org / 10.1080 / 00222939800770511","Moret P. 1989. Demembrement du genre Colpodes auctorum. I: Individualisation et definition des genres neotropicaux Dyscolus Dejean et Stenocnemion gen. nov. (Col. Caraboidea Platyninae). Bulletin de la Societe entomologique de France 93 (5 - 6): 133 - 148."]}

Dyscolus denigratus (Bates, 1891) Figs 45���50 Colpodes denigratus Bates, 1891: 17. Dyscolus palatus Moret, 1998: 18. syn. nov. Dyscolus (Dyscolus) denigratus ��� Moret 2005: 128. Dyscolus (Dyscolus) palatus ��� Moret 2005: 130. Dyscolus denigratus is a highly variable species, with a broad range of variation within each population regarding body size, form of the pronotum, length of the legs, and form of the apex of the male aedeagus (Figs 45���46, 48���50). The tree based on the COI sequences separates two clades (Fig. 2): one in the south-west part of the area of D. denigratus, represented in our dataset by the Cotopaxi and Pichincha populations, vs a north and north-east zone represented by the Cotacachi, Cayambe and Guaman�� populations. Additionally, three species-level units are recognized through BIN analysis: the first one embraces the populations of the south-western clade, including specimens identified as D. palatus. The northern clade is split into two BIN units: one for the Guaman�� population, and another for the Cotacachi and Cayambe populations. However, we were unable to find any morphological character that reliably reflects these divisions. According to the form of the apex of the median lobe, the Cotacachi population (Fig. 49) seems to be closer to the Pichincha population (Fig. 50) than to the Guaman�� one (Fig. 48), in contradiction with the molecular results. Obviously, the phylogeny of the denigratus complex cannot be resolved from the COI sequences alone. We thus take a conservative approach, with no taxonomic decision regarding the specific or subspecific status of the components of the denigratus complex, except for the synonymy of Dyscolus palatus with Dyscolus denigratus. Dyscolus palatus was described from specimens from the Pichincha, Atacazo and Coraz��n volcanoes in the Western Cordillera (Fig. 47). This taxon was then thought to be more closely related to D. altarensis (Bates, 1891) than to D. denigratus, due to its relatively broader and shorter body. The specimen of D. palatus included in our analysis (COI voucher PM359-01, BOLD sequence SUM168- 18) instead suggests that D. palatus and D. denigratus are conspecific. A re-examination of the type series of D. palatus and of large series of specimens recently collected on the Pichincha shows that the body dimensions of the two taxa partly overlap (Fig. 51). Owing to the existence of these transition forms, D. palatus cannot be maintained as a good species and we therefore treat it as a synonym of D. denigratus., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on page 43, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682, {"references":["Bates H. W. 1891. Coleoptera. In: Whymper E. (ed.) Supplementary Appendix to Travels amongst the Great Andes of the Equator: 7 - 39. John Murray, London.","Moret P. 1998. Les Dyscolus de la zone periglaciaire des Andes equatoriennes (Coleoptera, Harpalidae, Platyninae). Bulletin de la Societe entomologique de France 103 (1): 11 - 28.","Moret P. 2005. Los Coleopteros Carabidae del Paramo en los Andes del Ecuador: Sistematica, Ecologia y Biogeografia. Pontificia Universidad Catolica del Ecuador, Centro de Biodiversidad y Ambiente, Monografia 2, Quito."]}

Dyscolus incommunis Moret sp. nov. urn:lsid:zoobank.org:act: 696C59CA-E6A8-46CA-847F-CD9DA0167F60 Figs 9���10 Etymology Latin adjective meaning ���not together, not in common���; alluding to the fact that this species is not identical to D. aquator Moret sp. nov., despite the results of BIN analysis. Type material Holotype ECUADOR ��� ♂; Pichincha Province, Tandayapa, Bellavista Lodge, Waypoint 81; 0��0���56.6��� S, 78��40���49.1��� W; 2250 m a.s.l.; 2 Nov. 2015; P. Moret leg.; bromeliad fallen on the ground; QCAZ. Paratypes (2 ♂♂) ECUADOR ��� Pichincha Province ��� 1 ♂; same collection data as for holotype; leaf litter at night; COI voucher PM081-02, BOLD sequence SUM214-18; CPM ��� 1 ♂; Tandayapa, Bellavista Lodge; 0.0147�� S, 78.6833�� W; 2200 m a.s.l.; 9 Nov. 2010; D. Maddison leg.; COI voucher PM505, BOLD sequence SUM305-18; OSAC. Diagnostic description Habitus: Fig. 10. Wingless. Body length: 9.6���10.7 mm. Shiny piceous black, femora dark brown, the rest of the legs, antennae and mouthparts reddish brown. Elytral microsculpture transverse. Eyes moderately bulging, genae flat, mandibles long and acute. Pronotum convex, relatively big, elongate, narrowed backward but not cordiform; sides slightly sinuate posterad, hind angles obtuse and blunt; two pairs of lateral setae. Elytra elongate, humeri obliquely sloped, striae entire, finely impressed, with slight traces of punctation; third interval with three setae; subapical sinuation moderate; apical area triangle-shaped, rounded at apex. Last visible abdominal ventrite with one pair (♂) or two pairs (♀) of setae along its apical margin. Legs slender; fourth metatarsomere with or without subapical dorsolateral setae (one seta is present on the external lobe of the paratype), apical lobes parallel, the outer lobe 1.5 times longer than the inner lobe. Male genitalia: Fig. 9. Median lobe evenly arcuate, apex finely acute and slightly sinuate in lateral view, apex of the endophallus with a sclerotized structure formed of scales and denticles. Female genitalia: unstudied. Comparisons Although the BIN analysis suggests that D. aquator Moret sp. nov. and D. incommunis Moret sp. nov. are conspecific, the latter is differentiated from the former by important and stable morphological characters: sides of the pronotum less sinuate with obtuse hind angles; elytra broader and more convex, humeri not completely effaced, subapical sinuation weak, no apical tooth. The aedeagus��� median lobe is more arcuate, with a shorter apex and a sclerotized area differently shaped in the endophallus. Both species are sympatric in their type locality (Bellavista Lodge), but they live in different habitats. Habitat Montane forest on the Western slope of the Andes, at around 2250 m a.s.l. Unlike its closest relatives, this species has been collected on the forest ground, far from any stream, in the leaf litter and in fallen bromeliads. Geographic distribution Southern end of the Choc�� biogeographic region in northwestern Ecuador. Only known from the type locality, probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 15-16, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus donosoi Moret sp. nov. urn:lsid:zoobank.org:act: F31CFECF-7D04-4800-9559-EA234039B5DD Figs 15���16 Etymology Noun in the genitive case, dedicated to David A. Donoso, assistant professor at the Escuela Polit��cnica Nacional (Quito), specialist of the ecology of neotropical ants, who took part in the discovery of the holotype of this new species. Type material Holotype ECUADOR ��� ♀ (lacks the right foreleg and the right middle leg); Zamora-Chinchipe Province, Parque Nacional Podocarpus, Estaci��n El Colibr��; 3��59���16.1��� S, 79��5���39.0��� W; 2110 m a.s.l.; 18 Mar. 2015; P. Moret, C. Ruiz and D. Donoso leg.; COI voucher PM402-02, BOLD sequence SUM187-18; QCAZ. Paratypes (2 ♂♂) ECUADOR ��� 1 ♂ (lacks middle and hind legs); Zamora-Chinchipe Province, Parque Nacional Podocarpus, Fundaci��n Arcoiris; 3.988�� S, 79.093�� W; 2140 m a.s.l.; C. Ruiz and D. Mar��n leg.; COI voucher CR017, BOLD sequence GBNP 014-17; CISEC ��� 1 ♂ (lacks middle and hind legs); same collection data as for preceding; COI voucher CR018, BOLD sequence GBNP 015-17; CISEC. Diagnostic description Habitus: Fig. 15. Wingless. Body length: 11.8���13.9 mm. Body shiny black with a bluish, faintly iridescent lustre on the elytra; femora and tibiae nigropiceous, tarsi and palpi reddish brown; antennomeres 1���3 brunneopiceous, 4���11 reddish brown. Elytral microsculpture transverse. Head elongate, moderately convex, eyes relatively small, not protruding, genae slightly convex; mandibles very long, acute and arcuate at apex. Pronotum large, oval, completely rounded basally, with only one pair of lateral setae, lacking the basal pair; laterobasal depressions shallow. Elytra elongate-oval, convex; base arcuate; striae entire, broad and deeply impressed; intervals flat on disc, subconvex near apex; preapical sinuation obsolete. Third elytral interval without discal setae. Last visible abdominal ventrite. With one pair (♂) or two pairs (♀) of setae along its apical margin. Legs slender, fourth metatarsomere with one pair of subapical dorsolateral setae, apical lobes asymmetrical with a big outer lobe and no inner lobe. Male genitalia: Fig. 16. Median lobe arcuate with a long, spatulate apex, endophallus unarmed. Female genitalia: unstudied. Comparisons Owing to the unique combination of a rounded basal half of pronotum, absence of pronotal hind setae, and asetose third interval of the elytra, Dyscolus donosoi Moret sp. nov. has no close relatives among the described Ecuadorian Dyscolus. In our molecular analysis (Fig. 2) D. donosoi Moret sp. nov. forms a clade with D. saxatilis Moret, 1993, but the bootstrap support for this relationship is very weak. Dyscolus saxatilis is a p��ramo high-altitude species with black non-metallic integuments, broad head, short mandibles, and short and robust appendages. Habitat Montane forest on the Eastern slope of the Andes, at around 2100���2200 m a.s.l. Active at the beginning of the night (7���8.30 pm), on the trunk of a medium-sized tree, one meter above ground. Geographic distribution Only known from the type locality in Southern Ecuador, in the Parque Nacional Podocarpus. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 20-22, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus verecundior Moret sp. nov. urn:lsid:zoobank.org:act: DAB1A22E-25D1-438C-AE2E-25542AF7E50F Figs 54, 58 Dyscolus verecundus Moret, 1998: 20, pro parte. Dyscolus (Dyscolus) verecundus ��� Moret 2005: 142, pro parte. Etymology Comparative form of the Latin adjective ��� verecundus ���, which is the name of a closely related species. Type material Holotype ECUADOR ��� ♂; Pichincha Province, Volc��n Illiniza, ENE slope, Waypoint 136; 0.644404�� S, 78.703469�� W; 4335 m a.s.l.; 15 Jul. 2016; P. Moret and A. Barrag��n leg.; COI voucher in ethanol PM136-04, BOLD sequence SUM062-18; MNHN. Paratypes (2 ♂♂) ECUADOR ��� Pichincha Province ��� 1 ♂ (red printed label: ��� Dyscolus verecundus Moret, 1998, Paratype���); Volc��n Illiniza, NE slope; 4080 m a.s.l.; 7 Aug. 1985; P. Moret leg.; Polylepis forest; CPM ��� 1 ♂ (red printed label: ��� Dyscolus verecundus Moret, 1998, Paratype���); E slope of Volc��n Coraz��n; 4450 m a.s.l.; 24 Jan. 1986; P. Moret leg.; CPM. Diagnostic description Habitus: Fig. 54. Wingless. Body length: 10.8���11.0 mm. Body colour variable, from reddish brown to piceous black, the vertex and the sutural zone of the elytra reddish; legs, antennae and mouthparts reddish. Elytral microsculpture isodiametric, distinctly impressed. Head robust, convex, neck strongly constricted, eyes moderately bulging, genae almost flat in dorsal view; antennae short. Pronotum transverse, subquadrate; sides moderately arcuate distally, weakly sinuate in basal third; hind angles variable, obtuse or almost quadrate, but always blunt; two pairs of lateral setae. Elytra subparallel, disc almost depressed; humeri rounded but not effaced, subapical sinuation obsolete; striae thin, shallowly impressed; intervals flat; third elytral interval with 3���5 discal setae. Fourth metatarsomere triangular, with one pair of lateroapical setae and without subapical dorsolateral setae; apical lobes very short, the inner one not extending beyond the insertion point of the lateroapical seta; fifth metatarsomere asetose ventrally. Last visible abdominal ventrite of the male with one pair of setae along its apical margin (♂). Male genitalia: Fig. 58. Median lobe weakly arcuate; apical blade bluntly triangular in dorsal view; endophallus with two sclerotized squamose areas, ovoid in shape, median and subapical. Female genitalia: unknown. Comparisons This species closely resembles D. verecundus, now restricted to Mt Pichincha, its type locality. Dyscolus verecundior Moret sp. nov. differs from D. verecundus in a few external and genital characters: genae almost flat (convex in D. verecundus); antennae slightly thicker, the second antennomere of the male 1.41 to 1.49 times as long as wide (1.50 to 1.65 times in D. verecundus); fourth metatarsomere thinner, 1.16 to 1.20 times as long as wide (1.07 to 1.13 times in D. verecundus); apical blade of the median lobe thinner in lateral view (compare Figs 56���57 and 58); sclerotized areas of the endophallus bigger. Habitat Lower superp��ramo from 4080 to 4450 m a.s.l. Geographic distribution Microendemic species, restricted to the Illiniza and Coraz��n volcanoes in the Western Cordillera., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 47-48, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682, {"references":["Moret P. 1998. Les Dyscolus de la zone periglaciaire des Andes equatoriennes (Coleoptera, Harpalidae, Platyninae). Bulletin de la Societe entomologique de France 103 (1): 11 - 28.","Moret P. 2005. Los Coleopteros Carabidae del Paramo en los Andes del Ecuador: Sistematica, Ecologia y Biogeografia. Pontificia Universidad Catolica del Ecuador, Centro de Biodiversidad y Ambiente, Monografia 2, Quito."]}

Dyscolus crespoae Moret sp. nov. urn:lsid:zoobank.org:act: AEA12495-C2BD-4A65-AACF-C54DB6CAFBA7 Fig. 29 Etymology Noun in the genitive case, dedicated to Ver��nica Crespo, professor at the Pontificia Universidad Cat��lica del Ecuador, limnologist and specialist of the ecology of Andean tropical streams. Type material Holotype ECUADOR ��� ♀; Zamora-Chinchipe Province, Parque Nacional Podocarpus, Estaci��n El Colibr��; 3��59���16.1��� S, 79��5���39.0��� W; 2110 m a.s.l.; 18 Mar. 2015; P. Moret and C. Ruiz leg.; by night 19.30��� 21 pm; on the ground; QCAZ. Paratype ECUADOR ��� 1 ♀; same collection data as for holotype; COI voucher PM402-03, BOLD sequence SUM188-18; CPM. Diagnostic description Habitus: Fig. 29. Wingless. Body length: 10.6 mm (paratype) to 11.3 mm (holotype). Head and pronotum black, elytra nigropiceous; legs, antennae and mouthparts brownish to reddish brown. Dorsal integuments shiny, elytral microsculpture transverse. Head relatively small, elongate, eyes prominent, genae long, almost flat. Pronotum cordiform; sides markedly sinuate basally, hind angles obtuse and sharp; laterobasal impressions deep, lateral margins reflexed; two pairs of lateral setae. Elytra oval, convex; striae entire, shallowly impressed, distinctly punctate; intervals flat; preapical sinuation weakly marked. Third elytral interval without discal setae. Last visible abdominal ventrite with two pairs of setae along its apical margin. Legs moderately slender, all tarsi densely pubescent ventrally; metatarsomeres 1���3 bisulcate; external lobe of the fourth metatarsomere 2.5 times longer than inner lobe; fifth metatarsomere asetose ventrally. Male unknown. Female genitalia: unstudied. Comparisons This species belongs to the same clade as D. ravidus Moret sp. nov. Both have the same general aspect and share the character of an asetose third interval, but in D. crespoae Moret sp. nov. the dorsal surface is shiny with a transverse microsculpture on the elytra, the elytra are more convex, the hind angles of the pronotum more obtuse, the ventral pubescence of the tarsi denser, the apical lobes of the fourth metatarsomere more asymmetrical. Habitat Montane forest on the Eastern slope of the Andes, at around 2100 m a.s.l. Active at the beginning of the night (7.30���9 pm) on the surface of the leaf litter above ground. Geographic distribution Only known from the type locality in Southern Ecuador, in the Parque Nacional Podocarpus. Probably microendemic., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 30-32, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682

Dyscolus barragani Moret sp. nov. urn:lsid:zoobank.org:act: ACA0E4AD-2254-492D-9BD9-0107CDD395F6 Figs 43���44 Etymology Noun in the genitive case, dedicated to ��lvaro Barrag��n, associate professor of entomology at the Pontificia Universidad Cat��lica del Ecuador, co-organizer of the research project that led to the discovery of most of the new species described here. Type material Holotype ECUADOR ��� ♂; Chimborazo Province, Ayapungu, Cerro P��lpito, Waypoint 417; 2��18���16.2��� S, 78��34���33.4��� W; 4290���4320 m a.s.l.; 24 Mar. 15; P. Moret and A. Barrag��n leg.; QCAZ. Paratype ECUADOR ��� 1 ♂; Chimborazo Province, Ayapungu, Cerro P��lpito, Waypoint 416; 2��18���29.0��� S, 78��34���38.8��� W; 4180 m a.s.l.; 24 Mar. 15; P. Moret and A. Barrag��n leg.; COI voucher PM417-02, BOLD sequence SUM021-18; CPM. Diagnostic description Habitus: Fig. 43. Wingless. Body length: 8.9���9.2 mm. Body with a varying coloration, from reddish brown to piceous black, the sides of the pronotum and of the elytra lighter than the discal area; legs, antennae and mouthparts testaceous to reddish brown. Elytral microsculpture isodiametric, distinctly impressed. Head broad and convex, neck markedly constricted, eyes moderately bulging, genae oblique, almost flat in dorsal view; antennae moniliform, very short. Pronotum transverse, quadrate; lateroapical lobes small, not protruding; sides feebly arcuate distally, almost straight in basal third; hind angles bluntly obtuse, almost rounded; two pairs of lateral setae. Elytra subparallel, convex; preapical sinuation obsolete; striae fine and shallowly impressed, almost obsolete; intervals flat or subconvex; third elytral interval without discal setae. Legs short and robust; fourth metatarsomere triangular, with one pair of apical dorsolateral setae; apical lobes very short, the outer lobe slightly larger than the inner lobe; fifth metatarsomere asetose ventrally. Last visible abdominal ventrite of the male with one pair of setae along its apical margin. Male genitalia: Fig. 44. Median lobe weakly arcuate with a long apical blade, endophallus unarmed. Female genitalia: unknown. Comparisons This species closely resembles Dyscolus saxatilis Moret, 1993 in its general sturdy aspect, asetose third elytral interval, shallow striae, and in the form of the fourth metatarsomere. It differs from it by its smaller size (D. saxatilis: 10.0��� 10.8 mm), elytral microsculpture less granulose, eyes bigger, genae shorter and flat (convex in D. saxatilis), lateroapical lobes of the pronotum smaller. Our molecular analysis (Fig. 2) puts D. barragani Moret sp. nov. closer to D. hebeculus (Bates, 1891), with a very weak bootstrap support. Dyscolus hebeculus shares with D. barragani Moret sp. nov. an asetose third elytral interval, but otherwise the morphological differences are many. Dyscolus hebeculus has smaller eyes, longer antennae, a more elongate pronotum with more developed hind angles, elytral striae well impressed, and markedly asymmetrical lobes at the apex of the fourth metatarsomere. Habitat Humid superp��ramo, at around 4300 m a.s.l. Geographic distribution Microendemic species, restricted to the Ayapungu massif., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on pages 42-43, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682, {"references":["Bates H. W. 1891. Coleoptera. In: Whymper E. (ed.) Supplementary Appendix to Travels amongst the Great Andes of the Equator: 7 - 39. John Murray, London."]}

Genus Dyscolus Dejean, 1831 Dyscolus Dejean, 1831: 437. Type species: Dyscolus memnonius Dejean, 1831, by subsequent designation (Chaudoir 1878: 343). Hydrodyscolus Moret, 1996: 498. syn. nov. Dyscolus ��� Moret 1989: 143. The revision of the sequenced species led us to re-examine related material and to describe three new species of which we do not have DNA data, but which can be readily separated from the sequenced taxa (Dyscolus placitus Moret sp. nov., D. rivinus Moret sp. nov. and D. verecundissimus Moret sp. nov.). All the species under study share the following characters, which are, therefore, not repeatedly mentioned in the descriptions of the new species: labrum with six setae near apical margin; clypeus with two pairs of setae; two supraorbital setae each side; mentum with one pair of setae, submentum with two pairs of setae (except in the species of the hirsutus and verecundus groups, which may have additional submental setae); pronotum with two pairs of lateral setae, the distal one located near the widest point of pronotum, the basal one near or at the laterobasal angle (except D. donosoi Moret sp. nov. which lacks the basal seta); parascutellar setiferous pore present; metathoracic wings reduced and metepisternum very short (except in D. eleonorae Moret sp. nov.). The order in which these 25 new species are presented below is based on broad biogeographic and ecological groupings and on resemblances in external morphology, in order to make comparisons and identifications easier. The five resulting sections are: 1/ full-winged montane forest species (pp. 11���13); 2/ riparian wingless species of the montane forest (pp. 13���18); 3/ wingless, non-riparian species of the montane forest (pp. 18���36); 4/ riparian species of the p��ramo (pp. 36���42); 5/ non-riparian species of the p��ramo (pp. 42���51). All the new species are authored by P. Moret., Published as part of Moret, Pierre & Murienne, J��r��me, 2020, Integrative taxonomy of the genus Dyscolus (Coleoptera, Carabidae, Platynini) in Ecuadorian Andes, pp. 1-55 in European Journal of Taxonomy 646 on page 10, DOI: 10.5852/ejt.2020.646, http://zenodo.org/record/3829682, {"references":["Dejean P. F. 1831. Species general des Coleopteres de la collection de M. le comte Dejean. Tome cinquieme. Mequignon-Marvis, Paris. https: // doi. org / 10.5962 / bhl. title. 8863","Chaudoir M. de 1878. Revision des genres Onychopterygia, Dicranoncus et Colpodes. Annales de la Societe entomologique de France (5 e Serie) 8: 275 - 382.","Moret P. 1996. Mise au point taxinomique sur le genre Dyscolus Dejean et description de cinq especes nouvelles (Caraboidea, Harpalidae, Platynini). Bollettino del Museo regionale di Scienze naturali - Torino 14 (2): 491 - 503.","Moret P. 1989. Demembrement du genre Colpodes auctorum. I: Individualisation et definition des genres neotropicaux Dyscolus Dejean et Stenocnemion gen. nov. (Col. Caraboidea Platyninae). Bulletin de la Societe entomologique de France 93 (5 - 6): 133 - 148."]}

Grandcolas, Philippe, Legendre, Frédéric, Park, Yung Chul, Bellés, Xavier, Murienne, Jérôme, Pellens, Roseli (2005): The genus Cryptocercus in East Asia: distribution and new species (Insecta, Dictyoptera, Blattaria, Polyphagidae). Zoosystema 27 (4): 725-732, DOI: http://doi.org/10.5281/zenodo.4689649, {"references":["ASAHINA S. 1991. - Notes on two small collections of the Blattaria from China and Korea. Akitu 121: 1-5.","BEY- BIENKO G. 1935. - Descriptions of six new species of Palearctic Blattodea. Konowia 14: 117- 134.","BEY- BIENKO G. 1938. - On some new or interesting Asiatic Blattodea. Annals and Magazine of Natural History 1: 230-238.","BEY- BIENKO G. I. 1950. - [Fauna of the USSR. Insects. Blattodea s. Institute of Zoology, Academy of Sciences, Moscow, 342 p. (in Russian).","BRUGEROLLE G., SILVA- NETO I. D., PELLENS R. F GRANDCOLAS P. 2003. - Electron microscopic identification of the intestinal protozoan flagellates of the xylophagous cockroach Parasphaeria boleiriana from Brazil. Parasitology Research 90: 249-256.","CLARK J. W. , HOSSAIN S., BURNSIDE C. A. F KAMBHAMPATI S. 2001. - Coevolution between a cockroach and its bacterial endosymbiont: a biogeographical perspective. Proceedings of the Royal Society of London B 268: 393-398.","GADE G., GRANDCOLAS P. F KELLNER R. 1997. - Structural data on hypertrehalosaemic neuropeptides from Cryptocercus punctulatus and Therea petiveriana: how do they fit into the phylogeny of cockroachesx Proceedings of the Royal Society of London B 264: 763-768.","GRANDCOLAS P. 1994. - Phylogenetic systematics of the subfamily Polyphaginae, with the assignment of Cryptocercus Scudder, 1862 to this taxon (Blattaria, Blaberoidea, Polyphagidae). Systematic Entomology 19: 145-158.","GRANDCOLAS P. 1996. - The phylogeny of cockroach families: a cladistic appraisal of morphoanatomical data. Canadian Journal of Zoology 74: 508-527.","GRANDCOLAS P. 1997. - What did the ancestors of the woodroach Cryptocercus look likex A phylogenetic study of the origin of subsociality in the subfamily Polyphaginae (Dictyoptera, Blattaria), in GRANDCOLAS P. (ed.), The origin of biodiversity in insects: phylogenetic tests of evolutionary scenarios. Memoires du Museum national d'Histoire naturelle 173: 231-252.","GRANDCOLAS P. 1999. - Systematics, endosymbiosis, and biogeography of Cryptocercus clevelandi and C. punctulatus (Blattaria: Polyphagidae) from North America: a phylogenetic perspective. Annals of the Entomological Society of America 92: 285-291.","GRANDCOLAS P. 2000. - Cryptocercus matilei n. sp. du Sichuan de Chine (Dictyoptera, Blattaria, Polyphaginae). Revue francaise d'Entomologie (n.s.) 22: 223-226.","GRANDCOLAS P. F DELEPORTE P. 1996. - The origin of Protistan symbionts in termites and cockroaches: a phylogenetic analysis. Cladistics 12: 93-98.","GRANDCOLAS P. F D' HAESE C. 2004. - The origin of a \"true\" worker caste in termites: mapping the real world on the phylogenetic tree. Journal of Evolutionary Biology 17: 461-463.","GRANDCOLAS P., PARK Y. C., CHOE J. C., PIULACHS M. D., BELLES X., D' HAESE C., FARINE J. P. F BROSSUT R. 2001. - What does Cryptocercus kyebangensis n. sp. from Korea reveal about Cryptocercus evolutionx A study in morphology, molecular phylogeny and chemistry of tergal glands (Dictyoptera, Blattaria, Polyphagidae). Proceedings of the Academy of Natural Sciences of Philadelphia 151: 61-79.","KAMBHAMPATI S., CLARK J. W. F BROCK B. L. 2002. - Evolution of host- and habitat association in the wood-feeding cockroach. Biological Journal of the Linnean Society 75: 163-172.","MCKITTRICK F. A. 1964. - Evolutionary studies of cockroaches. Cornell University Agricultural Experiment Station, Memoir 389: 1-197.","NALEPA C. A. 2003. - Evolution in the genus Cryptocercus (Dictyoptera: Cryptocercidae): no evidence of differential adaptation to hosts or elevation. Biological Journal of the Linnean Society 80: 223-233.","NALEPA C. A., LI L. I., WEN- HUA L. F LAZELL J. 2001. - Rediscovery of the wood-eating cockroach Cryptocercus primarius (Dictyoptera: Cryptocercidae) in China, with notes on ecology and distribution. Acta Zootaxonomica Sinica 26: 184-190.","PARK Y. C. F CHOE J. C. 2003a. - Effects of parental care on offspring growth in the Korean wood-feeding cockroach, Cryptocercus kyebangensis. Journal of Ethology 21: 71-77.","PARK Y. C. F CHOE J. C. 2003b. - Territorial behavior of the Korean wood-feeding cockroach, Cryptocercus kyebangensis. Journal of Ethology 21: 79- 85.","PARK Y. C., GRANDCOLAS P. F CHOE J. C. 2002. - Colony composition, social behavior and some ecological characteristics of the Korean wood-feeding cockroach (Cryptocercus kyebangensis). Zoological Science 19: 1133-1139.","PARK Y. C., MAEKAWA K., MATSUMOTO T., SANTONI R. F CHOE J. C. 2004. - Molecular phylogeny and biogeography of the Korean woodroaches Cryptocercus spp. Molecular Phylogenetics and Evolution 30: 450-464.","PELLENS R., GRANDCOLAS P. F SILVA- NETO I. D. 2002. - A new and independently evolved case of xylophagy and the presence of intestinal flagellates in cockroaches: Parasphaeria boleiriana (Dictyoptera, Blaberidae, Zetoborinae) from the remnants of Brazilian Atlantic Forest. Canadian Journal of Zoology 80: 350-359.","WANG C. W. 1961. - The Forests of China, with a Survey of Grassland and Desert Vegetation. Maria Moors Cabot Foundation, Publication No. 5. Harvard University Press, Cambridge, Massachusetts, 313 p.","WILSON E. O. 1971. - The Insect Societies. The Belknap Press of the Harvard University Press, Cambridge, Massachusetts, 548 p.","YING T. S. 1983. - The floristic relationships of the temperate forest regions of China and the United States. Annals of the Missouri Botanical Garden 70: 597-604."]}

Cryptocercus parvus Grandcolas F Park, n. sp. (Figs 1B, D, H-J, N-P; 2C) TYPE MATERIAL. ��� Holotype *, allotype +, China, Heilongjiang, Laoheidingzi (1349 m) near Xingnong Linchang, 64 km WSW Hailin, SW Mudanjiang, X.1998, Yung Chul Park coll. (MNHN). Paratypes: same locality, 7 * *, 4 + + (MNHN); China, Heilongjiang, Laoheidingzi (1349 m) near Xingnong Linchang, 64 km WSW Hailin, SW Mudanjiang, X.1998, Yung Chul Park coll., 3 **, 3 ++ (in alcohol, NMNHK). TYPE LOCALITY. ��� Mudanjiang, China. OTHER MATERIAL EXAMINED. ��� China. Laotudingzi near Shuangfeng Linchang, 100 km WSW Hailin, 4 * *, 6 + +. ��� Tian-shan (= Tian Ling, or Mudanfeung), 1115 m, 32 km W Ningan, SW Mudanjiang, Yung Chul Park coll., 3 **, 2 ++ (MNHN). ETYMOLOGY. ��� The species is named according to its small size in the genus. MEASUREMENTS. ��� Body length: *, 16-18 mm; +, 15.5-17.5. Pronotum length: *, 5-5.2 mm; +, 4.2- 5.1 mm. DESCRIPTION Small species, short and wide, with punctuations on the dorsal surface.Pronotum(Fig.2C) with low but well delimited protuberances. Fore femora (Fig. 1D) with ventro-anterior margin with three to five spines, of roughly the same size, the more apical one generally farther from others especially in males. Male subgenital plate (Fig. 1B) with a medium space between styli, angularly curved. Male genitalia (Fig. 1 H-J) Sclerite L2d strong with a long and thin apex. L2v heavy, with a large sclerotized dorsal area and with an apical inner rounded part, and a ventral elongated and narrow part. L3 with a long and narrow apex. R3d with a blunt processus and an elongated basis with a rounded apex. R2 heavy but not wide. Female genitalia (Fig. 1 N-P) Basivalvulae (Fig. 1O) rounded with an inner margin irregularly sclerotized. Laterosternite IX (Fig. 1P) short, polygonal rather than triangular, with a short ventral apex. Spermatheca (Fig. 1N) with a large and rounded apical ampulla, and a curved basal ampulla on a short duct. DNA sequences The mitochondrial genes 12S and 16S have been partly sequenced for that species and are available on line in the EMBL database under the accession numbers AJ519680 and AJ519681 respectively., Published as part of Grandcolas, Philippe, Legendre, Fr��d��ric, Park, Yung Chul, Bell��s, Xavier, Murienne, J��r��me & Pellens, Roseli, 2005, The genus Cryptocercus in East Asia: distribution and new species (Insecta, Dictyoptera, Blattaria, Polyphagidae), pp. 725-732 in Zoosystema 27 (4) on pages 729-730, DOI: 10.5281/zenodo.4689649

Cryptocercus meridianus Grandcolas F Legendre, n. sp. (Figs 1A, E-G; 2B) TYPE MATERIAL. ��� Holotype *, China, Yunnan, Lijiang, cloud conifera forest, 3100 m, 23.VII.1993 (MNHN). TYPE LOCALITY. ��� Lijiang, Yunnan, China. ETYMOLOGY. ��� The species is named according to its very southern locality in China, near the border with Vietnam. MEASUREMENTS. ��� Body length: *, 24 mm. Pronotum length: *, 5.3 mm. DESCRIPTION Medium-sized species with a slender body. Body except pronotum smooth, not punctuated. Pronotum (Fig. 2B) with few and weak protuberances, except a protruding fore margin. Fore femora (Fig. 1E) with ventro-anterior margin with five spines not equally spaced and of roughly increasing size toward the apex. Male subgeni- t a l p l a t e (F i g. 1A) w i t h a w i d e a n d g e n t l y rounded margin between styli. Male genitalia (Fig. 1F, G) Sclerite L2d with an angular apex. Dorsal part of L2v also angular with a tooth protruding dorsoinnerly. Ventral part of L2v wide and strong. L1 with a rounded apex bearing one spine on the left and an expanded basis. R3d bulbose and slightly unsclerotized innerly, with the basal part sharp. DNA sequences The mitochondrial genes 12S and 16S have been partly sequenced for that species and are available on line in the EMBL database under the accession numbers AJ519679 and AJ519678 respectively., Published as part of Grandcolas, Philippe, Legendre, Fr��d��ric, Park, Yung Chul, Bell��s, Xavier, Murienne, J��r��me & Pellens, Roseli, 2005, The genus Cryptocercus in East Asia: distribution and new species (Insecta, Dictyoptera, Blattaria, Polyphagidae), pp. 725-732 in Zoosystema 27 (4) on pages 727-729, DOI: 10.5281/zenodo.4689649, {"references":["MCKITTRICK F. A. 1964. - Evolutionary studies of cockroaches. Cornell University Agricultural Experiment Station, Memoir 389: 1 - 197."]}