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The genus Anadia (family Gymnophthalmidae) consists of 19 species. It has remained almost taxonomically stable for decades, scarcely observed, in addition to being one of the less sampled gymnophthalmid genera with respect to molecular phylogenies. New Anadia species are discovered at a relatively low pace, and few specimens are found in the field, probably due to the arboreal habits of many low and mid elevation species. We describe here a new species of Anadia from the cloud forests of northwestern Colombia: the new species is easily diagnosed by the combination of shape and imbrication of dorsal scales, very large body size, the largest within its group, and large and non-overlapping number of longitudinal scale rows around midbody. We also tested the phylogenetic position of the recently described and geographically close A. antioquensis. A phylogenetic analysis based on four genomic regions recovered the new species as sister to A. buenaventura, whereas A. antioquensis was reassigned to the genus Riama. The new species is currently known from only three specimens, collected throughout eight years within less than 5 ha of the Mesenia-Paramillo Nature Reserve. Although its apparent rarity may be due to secretive habits, the species is provisionally declared vulnerable, while new information is available. To stimulate further research on this genus, we also compiled and present here the comparable information on the distribution and morphology of Anadia species. Altogether, the results stress the urge for a new review of the genus with the help of molecular data.

Anastelgis garciai Gauld, 1991 (Fig. 2E) Diagnosis. This species can be distinguished from all other Neotropical Anastelgis by the combination of the following characters: 1) body entirely reddish brown (propodeum more or less entirely reddish brown); 2) tegula reddish or infuscate; 3) epomia usually vestigial, generally just discernible as an indistinct rounded swelling; 4) submetapleural carina moderately strong, in lateral view clearly visible its entire length; 5) tergite III subquadrate to elongate, 1.1–1.6× longer than broad; 6) tergite II centrally virtually impunctate, at most with very scattered isolated punctures; 7) upper tooth of mandible blunt, very much broader than lower tooth (Gauld 1991). Distribution. Belize; Colombia (Arauca; Antioquia *); Costa Rica. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, El Oso, entomological net, N 5°29′54.5″, E 75°54′35.7″, 2.500 msnm, 09.VIII.2020 (J. Jaramillo leg.), 1♀, UNIANDES; idem, but Mcpio [=Municipio] Jardín, sweep net, N 5°29′, E 75°54′, 25.III.2020 (J. Jaramillo leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 65, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Gauld, I. D. (1991) The Ichneumonidae of Costa Rica, 1. Memoirs of the American Entomological Institute, 47, 1 - 589."]}

Epirhyssa fusca Gómez & Sääksjärvi, 2015 (Fig. 5B) Diagnosis. According to Gómez et al. (2015), this species can be distinguished from the other neotropical Epirhyssa by the combination of the following character states: 1) body primarily dark brownish with some yellowish parts; 2) mesoscutum with transverse rugae rounded (not sharp); 3) body finely punctate with abundant setae; 4) tergite I stout and with wrinkles, about 1.6× as long as posteriorly broad; 5) propodeum very short and finely granulate with long setae. Distribution. Colombia * and Peru. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Malaise trap 4, N 5°29′55.5″, E 75°54′22.9″, 2623 msnm, I.2021 (L. Mazariegos leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 69, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Gomez, I. C., Saaksjarvi, I. E., Puhakka, L., Castillo, C. & Bordera, S. (2015) The Peruvian Amazonian species of Epirhyssa Cresson (Hymenoptera: Ichneumonidae: Rhyssinae), with notes on tropical species richness. Zootaxa, 3937 (2), 311 - 336. https: // doi. org / 10.11646 / zootaxa. 3937.2.4"]}

Pimpla pyramis (Porter, 1970) (Fig. 4D) Diagnosis. This species can be distinguished from the other Colombian Pimpla by the combination of the following character states: 1) tergite I in profile with high, more or less obscure punctures; 2) wings hyaline with a slight to definite yellowish brown mark; 3) mesosoma shining black with postscutellum white, legs and fore coxae entirely black, metasoma black and tergites with white stripe on the back; 4) ovipositor cylindrical, dorsal valve gently convex on the tip; 5) apical margin of clypeus deeply emarginated; 6) gena 0.7–0.8× as long as basal as eye in lateral view; 7) metasoma with laterotergite V narrow, about 0.4× as broad as long (Porter 1970; Pádua et al. 2019). Distribution. Bolivia; Colombia (Antioquia *, Cundinamarca); Venezuela. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Malaise trap 6, N 5°29′54.5″, E 75°54′35.7″, 2797 msnm, VII.2020 (L. Mazariegos leg.), 1♀, UNIANDES; Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, sweep net, N 5°29′, E 75°53′, 06.I.2020 (J. Jaramillo leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 68, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Porter, C. C. (1970) A Revision of the South American Species of Coccygomimus (Hymenoptera, Ichneumonidae). Studia Entomologica, 13, 1 - 192.","Padua, D. G., Araujo, R. O. & Mazariegos, L. A. (2019) Pimpla Fabricius (Hymenoptera: Ichneumonidae: Pimplinae) from Colombia. Zootaxa, 4683 (3), 439 - 446. https: // doi. org / 10.11646 / zootaxa. 4683.3.8"]}

Dolichomitus mariajosae Araujo & Pádua, 2020 (Fig. 3A–C) Diagnosis. See Araujo et al. (2020) Male description (Fig. 3B). Lacking tergites III+. Similar to female in structure and coloration, except face with central part blackish; fore wing hyaline without a black spot; tergite I entirely yellow; legs with lighter black markings than those of females; face approx. 1.15× as width as high; posterior ocelli separated from eyes by 1.0× its maximum diameter; fore wing 7.0 mm; mid coxa with distinct basal and central prominences which are separated by a weak deep concavity (Fig. 3C); tarsal claw simple; metasoma slender, tergite I about 1.7× as long as posterior width; tergite II about 1.0× as long as posteriorly width. Distribution. Colombia (Antioquia). Material examined. COLOMBIA, Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, La Lucrecia, entomological net, N 5°29′54.5″, E 75°54′35.7″, 22.I.2020 (J. Jaramillo leg.), 1♀, UNIANDES; idem, but Mcpio [=Municipio] Jardín, sweep net, N 5°29′, E 75°54′, 22.IV.2020 (J. Jaramillo leg.), 1♀, UNIANDES; idem, but Olinguito, entomological net, [5°31′2.99″N, 75°51′39.33″W], 20.VI.2020 (J. Jaramillo leg.), 1♁ [metasoma without tergite III+], UNIANDES.

Forrestopius auguratricis Alvarado & Palacio, 2021 (Fig. 2C) Diagnosis. This species can be distinguished from all other Forrestopius by the combination of the following characters: 1) body black, with scape, dorso-posterior corner of propodeum yellowish brown, and fore and mid legs predominantly yellowish testaceous, and hind leg yellowish testaceous, coxa and basal half of femur black, distal 1/3 of tibia light brown; 2) mandible with two teeth; 3) antenna with 16–19 flagellomeres; 4) face and clypeus forming a continuous surface; 5) pronotum with a single wrinkle arising from pronotal pit with its upper end reaching about one third to half of way to anterior margin; 6) propodeum with lateromedian and lateral longitudinal carinae (Alvarado & Palacio 2021). Distribution. Colombia (Risaralda; Coquimbo*); Ecuador; Peru; Venezuela. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Malaise trap 3, N 5°29′55.5″, E 75°54′14.0″, 2501 msnm, IX.2020 (L. Mazariegos leg.), 4♀♀, UNIANDES; idem, but Malaise trap 4, N 5°29′55.5″, E 75°54′22.9″, 2623 msnm, II–III.2021, 1♀, UNIANDES; idem, but Malaise trap 5, N 5°29′54.3″, E 75°54′28.7″, 2707 msnm, I.2021, 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on pages 63-64, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Alvarado, M. & Palacio, E. (2021) Forrestopius Gauld & Sithole, 2002 (Hymenoptera: Ichneumonidae: Metopiinae) in South America. Zootaxa, 5040 (2), 265 - 282. https: // doi. org / 10.11646 / zootaxa. 5040.2.6"]}

Aphanistes silviae Alvarado, 2018 (Fig. 2A) Diagnosis. According to Alvarado (2018), this species can be distinguished from all other nocturnal Aphanistes Förster, 1869 by the combination of the following characters: 1) ovipositor with a dorsal notch and not pre-apically swollen; 2) body predominantly reddish-brown; 3) hind tarsomeres II–IV whitish cream. Distribution. Colombia * and Peru. Material examined. COLOMBIA, Mcpio [= Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, [Olinguito], Malaise trap GG [= Gressitt & Gressitt model], N 5°31′03″, E 75°51′50″, 2400 msnm, IX.2020 (L. Mazariegos leg.) 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 63, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Alvarado, M. (2018) Nocturnal Aphanistes (Hymenoptera: Ichneumonidae: Anomaloninae) in the Neotropical region, with the description of five new species. Zootaxa, 4369 (2), 221 - 236. https: // doi. org / 10.11646 / zootaxa. 4369.2.4"]}

Dolichomitus orejuelai Araujo & Pádua, 2020 (Fig. 3G–I) Diagnosis. See Araujo et al. (2020) Male description (Fig. 3H). Similar to female in structure and coloration, except mesosoma entirely blackish; mid coxa whitish and hind coxa and trochanters blackish; metasoma with tergite I black and tergites II+ reddish orange with black narrow bands; Body 16 mm; fore wing 12 mm; face about 1.1× as width as high; posterior ocelli separated from eyes by 1.0× its own maximum diameter; mid coxa with distinct basal, central, and apical small prominences which are separated by the weak deep concavities (Fig. 3I); tarsal claw simple; metasoma slender, tergite I about 1.9× as long as posterior width; tergite II about 1.7× as long as posterior width; paramere with apex large and weakly rounded. Distribution. Colombia (Antioquia). Material examined. COLOMBIA, Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, sweep net, N 5°29′, E75°54′, 11.VI.2020 (J. Jaramillo leg.), 1♁,UNIANDES; idem, but 14.IV.2020, 1♀,UNIANDES; idem, but 06.I.2020, 1♀, UNIANDES; Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, El Oso, entomological net, N 5°29′54.5″, E 75°54′35.7″, IX.2020 (J. Jaramillo leg.), 3♀♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 66, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Araujo, R. O., Padua, D. G., Jaramillo, J. & Mazariegos, L. A. (2020) Five new species of Dolichomitus Smith from the tropical Andes, with a key for the South American species (Hymenoptera, Ichneumonidae, Pimplinae). ZooKeys, 937, 89 - 113. https: // doi. org / 10.3897 / zookeys. 937.51361"]}

Neotheronia lineata (Fabricius, 1804) (Fig. 4A) Diagnosis. According to Gauld (1991), this species can be distinguished from all other Colombian Neotheronia by the combination of the following characters: 1) body yellow with a black stripe joining the lateral ocellus and eyes, a black area around ocelli and black stripe extending back to form a broad band on occiput, and black marks on anterior margin and a medial stripe running from transverse carina to the posterior margin in propodeum; 2) fore wing with a black spot in the apex (sometimes very weakly); 3) occipital carina with distinct notch mediodorsally; 4) pronotum with the upper end of epomia quite strong (about 0.5× the length of the basal mandibular width); 5) lateral longitudinal carina in front of transverse carina present in propodeum; 6) ovipositor 1.4–1.5× the length of hind tibia. Distribution. Argentina; Bolivia; Brazil; Colombia (Antioquia *); Cuba; Ecuador; Guyana; Mexico; Paraguay; Peru; Suriname; Trinidad & Tobago; Uruguay. Krieger (1905) registered N. lineata to Colombia, but did not mention the specific place (department o municipality) in which specimens were collected. Material examined. COLOMBIA, Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve (1800–3000 m), entomological net, N 5°29′45.8′′, E 75°53′21.3′′, VIII.2019 (L. Mazariegos leg.), 1♀ and 2♁♁, UNIANDES; idem, but XI.2018, 1♁, UNIANDES; idem, but III.2019, 1♁, UNIANDES; idem, but Enrielado, 11.III.2020, 1♀, UNIANDES; Mcpio [= Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, El Rodano, entomological net, 31.I.2020 (J. Jaramillo leg.), 1♁, UNIANDES; idem, but Cristalina, 23. VI.2020, 1♁, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on pages 66-67, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Gauld, I. D. (1991) The Ichneumonidae of Costa Rica, 1. Memoirs of the American Entomological Institute, 47, 1 - 589.","Krieger, R. (1905) Ueber die Ichneumonidengattung Neotheronia Krieger. (Hym.). Zeitschrift f ¸ r Systematische Hymenopterologie und Dipterologie, 5, 286 - 338."]}

Pimpla punicepis Cresson, 1874 (Fig. 4C) Diagnosis. This species can be distinguished from the other Colombian Pimpla by the combination of the following character states: 1) metasoma with laterotergites II–V broad, more than 0.5× as broad as long; 2) malar space wide, 1.0× longer than basal mandibular width (0.8–0.9× in male); 3) female with the posterolateral corner of mesopleuron usually finely punctate; 4) body black with legs reddish brown; 5) male with a central white band on hind tibia (Gauld 1991; Díaz 2000). Distribution. Chile; Colombia (Antioquia *, Boyacá, Magdalena, Nariño); Costa Rica; Ecuador; Guatemala; Honduras; Mexico; Peru; USA; Venezuela. Material examined. COLOMBIA, Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve (1800–3000 m), entomological net, N 5°29′45.8′′, E 75°53′21.3′′, VIII.2019 (L. Mazariegos leg.), 2♀♀, UNIANDES; idem but IV.2019, 4♀♀, UNIANDES; idem, but sweep net, N 5°29′, E 75°53′, 25.III.2020 (J. Jamarillo leg.), 1♀, UNIANDES; idem, but La Lucrecia, entomological net, N 5°29′54.5′′, E 75°54′35.7′′, IX.2020 (J. Jamarillo leg.), 2♁♁, UNIANDES; Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Mesenia, entomological net, N 5°29′54.5″, E 75°54′35.7″, 19.IX.2020 (J. Jamarillo leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on pages 67-68, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Gauld, I. D. (1991) The Ichneumonidae of Costa Rica, 1. Memoirs of the American Entomological Institute, 47, 1 - 589.","Diaz, F. A. (2000) The Venezuelan Species of Pimpla (Hymenoptera: Ichneumonidae). Journal of Hymenoptera Research, 9 (2), 246 - 253."]}

Pimpla albomarginata Cameron, 1886 (Fig. 4B) Diagnosis. This species can be distinguished from the other Colombian Pimpla by the combination of the following character states: 1) body reddish brown with white marks, except with head, mesoscutum, postscutellum, metasoma (with white bands) black, fore coxa with black mark; 2) wings hyaline; 3) clypeus with apex deeply bilobed; 4) malar space wide, about 0.7–0.8× longer than basal mandibular width (Rs more or less straight, and 1 cu-a slightly postfurcal to the base of M&RS; 7) propodeum with conspicuous posterolateral tubercles; 8) laterotergite V more than 0.5× as broad as long; 9) tergite I of female short and broad, in profile strongly convex in profile with moderately high blunt hump; 10) sternite I with strongly produced swelling; 11) apex of ovipositor with dorsal valve lacking teeth apically (Gauld 1991). Distribution. Colombia *; Costa Rica; Mexico; Panama; Uruguay; Venezuela. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Malaise trap 4, N 5°29′55.2″, E 75°54′22.9″, 2623 msnm, VII.2020 (L. Mazariegos leg.), 2♀♀, UNIANDES; idem, but Malaise trap 3, N 5°29′55.5″, E 75°54′14.0″, 2501 msnm, IX.2021, 1♀, UNIANDES; idem, but VIII.2020, 1♀, UNIANDES; Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve (1800–3000 m), entomological net, N 5°29′45.8′′, E 75°53′21.3′′, VIII.2019 (L. Mazariegos leg.), 2♀♀, UNIANDES; idem, but [Olinguito], Malaise trap GG [= Gressitt & Gressitt model], N 5°31′03″, E 75°51′50″, 2400 msnm, X–XI.2020, 1♀, UNIANDES; Mcpio [=Municipio] Mistrató, Risaralda, Mesenia-Paramillo Nature Reserve, sweep net, 27.VI.2020 (J. Jaramillo leg.), 1♀, UNIANDES. Comments. Pádua et. al. (2019) recorded P. albomarginata for first time in Colombia. However, the specimens analyzed by these authors were a misidentification. According to Dr. Edgar Palacio (pers. comm. with ROA), that species is a new species in the albomarginata group (sensu Porter 1970) which he will describe later. So, P. albormarginata is recorded for the first time in Colombia in this work., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 67, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Gauld, I. D. (1991) The Ichneumonidae of Costa Rica, 1. Memoirs of the American Entomological Institute, 47, 1 - 589.","Porter, C. C. (1970) A Revision of the South American Species of Coccygomimus (Hymenoptera, Ichneumonidae). Studia Entomologica, 13, 1 - 192."]}

Ganodes mexicanus Díaz, 2008 (Fig. 5A) Diagnosis. This species can be distinguished from the other Ganodes by the combination of the following character states: 1) face white with a narrow longitudinal black stripe; 2) posterior ocellus separated from eye by 1.5× its maximum diameter; 3) metapleuron sparsely and finely punctate; 4) hind coxa and femur white with longitudinal black stripes; 5) metasoma black with lateral and posterior margins of tergite I and posterior margin of remaining tergites all broadly white; 6) ovipositor 2.1× length of the hind tibia (Díaz 2008). Distribution. Colombia * and Mexico. Material examined. COLOMBIA, Mcpio [= Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, [Olinguito], Malaise trap GG [= Gressitt & Gressitt model], N 5°31′03″, E 75°51′50″, 2400 msnm, VIII.2020 (L. Mazariegos leg.) 1♀, UNIANDES; Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Cristalina, entomological net, 23.VI.2020 (J. Jaramillo leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 69, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Diaz, F. A. (2008) Five New Species of the Neotropical Genus Ganodes Townes (Hymenoptera: Ichneumonidae). Neotropical Entomology, 37 (6), 668 - 673."]}

Acrotaphus pseudoamazonicus Pádua & Sääksjärvi, 2020 (Fig. 2D) Diagnosis. According to the New World review of Acrotaphus by Pádua et al. (2020), this species can be distinguished from all other species by the combination of the following characters: 1) body orange except head, posterolateral bands in the tergites II–IV, tergite V with distal half and VI+, hind leg black (in generally); 2) margin of gena behind eyes flat in dorsal view; 3) margin of gena 0.35–0.5× (0.5–0.6 in male) length of eye in dorsal view; 4) posterior ocelli separated from eyes by 0.3–0.5× its diameter in dorsal view; 5) ovipositor robust, 1.1–1.5× as long as hind tibia.. Distribution. Brazil; Colombia (Putumayo; Chocó; Antioquia *); Ecuador; French Guiana; Panama; Peru. Material examined. COLOMBIA, Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, El Rodano, entomological net, N 5°29′54.5″, E 75°54′35.7″, 05.I.2020 (J. Jaramillo leg.), 1♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 64, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Padua, D. G., Saaksjarvi, I. E., Monteiro, R. F. & Oliveira, M. L. (2020) Review of the New World genus Acrotaphus Townes, 1960 (Hymenoptera: Ichneumonidae: Pimplinae), with descriptions of fifteen new species. Zootaxa, 4719 (1), 1 - 62. https: // doi. org / 10.11646 / zootaxa. 4719.1.1"]}

Pádua, Diego G., Fernandes, Daniell R.R., Araujo, Rodrigo O., Mazariegos, Luis A. (2023): Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes. Zootaxa 5244 (1): 61-70, DOI: 10.11646/zootaxa.5244.1.5, URL: http://dx.doi.org/10.11646/zootaxa.5244.1.5

Woldstedius paulus Dasch, 1964 (Fig. 2B) Diagnosis. This species can be distinguished from all other Woldstedius by the combination of the following characters: 1) hind wing with CU absent; 2) external face of the hind tibia without setae; 3) mesopleurum with a longitudinal stripe; 4) fore and mid legs mostly yellow, except mid tibia and tarsus dusky dorsally, hind coxa yellow (except posterior part fuscous) (Dasch 1964). Distribution. Colombia (Antioquia: Medellín; Jardín*; Andes*); Costa Rica; Ecuador; Mexico. Material examined. COLOMBIA, Mcpio [=Municipio] Jardín, Antioquia, Mesenia-Paramillo Nature Reserve, [Olinguito], Malaise trap GG [= Gressitt & Gressitt model], N 5°31′03″, E 75°51′50″, 2400 msnm, I.2021 (L. Mazariegos leg.) 1♀, UNIANDES; Mcpio [=Municipio] Andes, Antioquia, Mesenia-Paramillo Nature Reserve, Malaise trap 3, N 5°29′55.5″, E 75°54′14.0″, 2501 msnm, VIII.2020 (L. Mazariegos leg.), 2♀♀, UNIANDES., Published as part of Pádua, Diego G., Fernandes, Daniell R. R., Araujo, Rodrigo O. & Mazariegos, Luis A., 2023, Filling gaps in the knowledge of the Colombian Darwin wasps (Hymenoptera: Ichneumonidae): new records, description of unknown males, and taxonomical notes, pp. 61-70 in Zootaxa 5244 (1) on page 63, DOI: 10.11646/zootaxa.5244.1.5, http://zenodo.org/record/7645791, {"references":["Dasch, C. E. (1964) The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae). Contributions of the American Entomological Institute, 1 (1), 1 - 78."]}

This research is focused to understand how the occupancy of the Andean tiger cat is influenced by different drivers acting at the microhabitat at landscape scales in protected cloud forests from northwestern Colombia.

Anadia hollandi sp. nov. Holotype. MHUA-R 13725 (Fig. 3). Male collected at the field station of the Mesenia-Paramillo Nature Reserve (5.496º N, 75.889º W), at 2170 m asl, Vereda La Mesenia, about 14 km south of the municipality of Jardín (Department of Antioquia), Colombia. The specimen was collected by Diana Rendón on September 1 st, 2011. Paratypes. MHUA-R 13726 (field code GECOH3499), and MHUA-R 13727 (field code LSB409) (Fig. 3). Two presumably adult males collected within 5 km from the field station, on January 25 th 2018 by Ubiel Rendón, and on June 26 th 2019 by Carlos Rendón, respectively. Etymology. The specific epithet hollandi is a patronym in honour of Jim Holland, American olympic skier who enjoys the natural world and the challenges of outdoor exploration, for his generous contribution to the conservation of cloud forests in Colombia. Definition and diagnosis. The largest species of Anadia known to date in the ocellata group (Fig. 4), reaching 108 mm of SVL and 351 mm of TL. It can be diagnosed by the following combination of morphological characters: (1) subhexagonal smooth dorsal scales; (2) two prefrontal scales in broad (between 20–50% of scale size) contact; (3) divided nasal scales; (4) three supraocular scales; (5) 0–1 inserted scales between superciliary and supraocular scales; (6) five superciliary scales; (7) lower palpebrals without pigment; (8) five subocular scales; (9) 6–8 supralabial scales; (10) 5–6 infralabial scales; (11) seven postparietal scales; (12) first two pairs of genial scales in contact; (13) 43–45 transverse rows of dorsal scales; (14) 33–36 transverse rows of ventral scales; (15) 1.19–1.36 transverse rows of dorsal scales for each ventral row; (16) quadrangular ventral scales; (17) 41–43 scale longitudinal rows around mid body; (18) 10–12 supradigital scales on fourth toe; (19) 15–18 lamellae under fourth toe; (20) 10–12 femoral pores in males (no available females); (21) 7–9 large black ocelli with white centers per body side; (22) belly cream with scattered dark spots. Anadia hollandi sp. nov. can be easily distinguished from all other Anadia species by a combination of the shape and imbrication of dorsal scales, and the number of scale rows around midbody. In particular [A. hollandi sp. nov. in brackets], all members of the eastern group, and therefore of the bitaeniata, steyeri and marmorata species groups have longer, rectangular and juxtaposed [versus shorter, subhexagonal and imbricate] dorsal scales; within the western group, A. bogotensis has considerable imbrication of dorsal and ventral scales [versus subimbricate dorsal scales]; A. bogotensis in the western group and all other members of the ocellata group have a much lower number of longitudinal scale rows around mid-body: 24–32 in A. bogotensis and 27–32 to 30–37 [versus 41–43] (Fig. 4). In addition, A. hollandi is the species with the largest body size [89.1–109.0 mm SVL] among the known species of the western group: 40.0 – 61.4 mm SVL in all other species (Fig. 4). Description of the holotype (Figs 3,5). A male, 89.1 mm of snout-vent length (SVL), 210.7 mm of tail length, and 299.8 mm of total length. Head length 20.9 mm, head width 13.1 mm, and snout length 17.2 mm. Head scales smooth, glossy, and juxtaposed. Rostral wider than high, mostly in contact with frontonasal, less with the first supralabial, and barely with the anterior division of the nasal. Frontonasal pentagonal, not elongated, and with a slightly rounded anterior edge, contacting laterally the divided nasals, and posteriorly the prefrontals. No contact between frontal and loreals. Prefrontals in mid contact, about 30% of the scale length, hexagonal, also contacting frontonasal, the posterior division of the nasal, the loreal, the presupraocular, and the frontal. Frontal pentagonal, slightly wider on the anterior edges, about the same size as the frontonasal, mostly in contact with prefrontals, supraoculars, and the frontoparietals; barely in contact with presupraoculars. Frontoparietal two, subhexagonal, contacting laterally the second and third pair of supraoculars, and posteriorly the parietals and the interparietal. Interparietal hexagonal, about twice as long as wide, contacting laterally the two pentagonal and larger parietals; parietals and interparietal contact posteriorly a row of seven uneven postparietals, also considered the first transversal row of dorsal scales. Supraoculars three, the anterior larger than the other two, in contact laterally with five superciliary scales. Divided nasal scales, in contact ventrally with first and second supralabials, posteriorly with loreal, and dorsally with dorsal and prefrontal scales. Nostril rounded, placed mostly within the anterior division of the nasal. Loreal pentagonal, in contact dorsally with prefrontal, posteriorly with presuperciliary and frenocular, and ventrally with the second supralabial. Frenocular contacting dorsally the presuperciliary, posteriorly the first subocular, and ventrally the second and third supralabials. Presuperciliary roughly pentagonal, in contact posteriorly with first superciliary and first ciliary, and expanded onto dorsal surface of head, contacting dorsally a small triangular presupraocular. Superciliary five, much larger at the anterior and posterior extremes, and in contact ventrally with ciliary. Six ciliary scales. Lower palpebrals translucent and unpigmented. Suboculars five, polygonal, first contacting posterior end of third supralabial, second contacting both third and fourth supralabials, third contacting fourth supralabial, fourth contacting both fourth and fifth supralabial, and fifth contacting fifth and marginally sixth supralabial. Postoculars three, polygonal. Supralabials seven, third longest. Mental wider than long, and in contact posteriorly with the first two pairs of infralabials and the postmental. Infralabials five, fifth the longest. Postmental single, pentagonal, wider than long, in contact posteriorly with the first pair of genials. Three pairs of genials, the anterior two pairs in broad medial contact. The first pair in direct contact with second and mostly third infralabials. The second pair of genials in contact with the third and the fourth infralabials. Third pair of genials in contact with fourth and fifth infralabials. Transverse gular scales in 14-15 rows between the posterior edge of the third pair of genials and the collar fold. Dorsal scales sub-hexagonal in shape, twice longer than wide, with straight lateral edges, - smooth and glossy. Dorsal scales arranged in 43 transverse rows. Thirty-one transverse dorsal scales between the posterior edge of forelimbs and the anterior edge of hindlimbs. Lateral scales similar to dorsal scales, smaller near the insertions of limbs. More rows (1.30 times) of dorsal transverse scales for each ventral row. First transverse row of pentagonal ventral scales -. From that point, all transverse ventral scales are roughly quadrangular, smooth and juxtaposed. Ventral scales 1.3–1.5 times longer than wide, and 1.2 times longer and 1.2 times wider than dorsal scales. Ventral scales arranged in 33 transverse rows. Scale rows around the mid body 41. Six posterior cloacal plate scales. Tail complete but the very tip, cylindrical, with scales rectangular, smooth, juxtaposed and arranged clearly in transverse rows. Limbs pentadactyl and digits clawed. Anterodorsal brachial scales polygonal, subequal in size, imbricate and smooth. Posteroventral brachial scales rounded, uneven in size, smooth and imbricate. Dorsal and anterior antebrachial scales rounded, subequal in size, smooth, and subimbricate. Scales on dorsal surfaceof manus polygonal, smooth and imbricate. Scales on manus palmar surface ovoid, juxtaposed, slightly smooth and smaller than dorsals. Thenar scales three. Scales on dorsal surface of fingers smooth, quadrangular, imbricate, overhanging supradigital lamellae, three on finger I, six on II, nine on III, nine on IV, and six on V. Subdigital lamellae three on finger I, eight on II, 11 on III, 14 on IV, and nine on V. Dorsal and anterior scales of thigh large, polygonal and juxtaposed. Posterior thigh scales small, rounded and slightly imbricate. Ventral thigh scales large, rounded and imbricate. Seven and eight femoral pores per leg, and no scales between each pair of femoral pores. Dorsal and anterodorsal crus scales polygonal, smooth and juxtaposed. Ventral and posterior crus scales rounded, small and imbricate. Scales on dorsal surface of toes rounded, smooth, varying in size. Supradigital lamellae four on toe I, seven on II, 10 on III, 10 on IV, and 10 on V. Subdigital lamellae single, five on toe I, nine on II, 13 on III, 12 on IV, and 12 on V. Coloration (Figs. 3, 5). In life, the holotype is light brown with irregular dark brown marks throughout trunk and near tail insertion. The dark marks become roughly rounded and connected throughout the rest of the dorsal tail. The paratypes lack dark brown marks and tail patterning. Venter is cream white, with few dark brown to black speckles. Near the head, the cream white coloration suffuses with light brown towards the supralabial scales. Each body flank bears a longitudinal row of 7–9 black ocelli with white centers. Limbs are dorsally light brown and ventrally cream white with few dark speckles. In specimens preserved in ethanol, light brown becomes darker, dark spots and speckles remain visible, and ventral cream white becomes dark grey. Hemipenes (Fig. 6). We extracted the hemipenes of both paratypes. They are bilobed with a similar length of truncus; the lobes are rounded with a reticulated surface and with the presence of small lateral calyces towards the sulcate side of the hemipene. The asulcated side exhibits several folds: at the base a horizontal fold, and towards the upper part of the truncus, laminar folds arranged in an inverted “V” shape, which reaches the middle of the truncus without touching the sulcus spermaticus. Natural history. All individuals were found near the cloud forest edge, one of them apparently basking on the roof of the Reserve’s biological station, and a second within an open area of the house. The third was spotted coming out a large arboreal bromeliad, at about 2 m above the ground level. In captivity, they often adopted a hanging posture, using their long tails and posterior limbs to hang upside down from small branches or sticks (Fig. 5). Near all species of Anadia are known or suspected to be arboreal, dwelling within bromeliad plants or the leaf litter accumulated between the large tree branches. Exceptions are the high-altitude species (e.g., A. bogotensis and A. brevifrontalis), which are mostly found on the ground, under rocks and logs, or within rosette plants,. Conservation status. The new species is only known from the edge of the Mesenia-Paramillo nature reserve and the specimens were collected within 5 ha. As mentioned above, the lack of sightings despite intensive search at the reserve may be due to the species’ arboreal habitat. Nonetheless, no specimens of this species were found in the large herpetological collections at the Universidad de Antioquia (Medellín, Colombia) and the Instituto de Ciencias Naturales of the Universidad Nacional de Colombia (Bogotá, Colombia). Due to the lack of information on its distribution, the relatively small area of the nature reserve, and the vulnerability of forests within this elevational belt, we suggest assigning A. hollandi sp. nov. to the category Vulnerable (VU: B1a, biii; IUCN, 2001), until new information is available., Published as part of Amézquita, Adolfo, Daza, Juan Manuel, Contreras, Leidy Alejandra Barragán, Orejuela, Catalina, Barrientos, Lucas Santiago & Mazariegos, Luis A., 2022, One more and one less: a new species of large bromelicolous lizard (Gymnophthalmidae: Anadia) from the Andean cloud forests of northwestern Colombia and the phylogenetic status of Anadia antioquensis, pp. 217-238 in Zootaxa 5150 (2) on pages 221-225, DOI: 10.11646/zootaxa.5150.2.3, http://zenodo.org/record/6621199, {"references":["Oftedal, O. T. (1974) A revision of the genus Anadia (Sauria, Teiidae). Arquivos de Zoologia, 25, 203 - 265. https: // doi. org / 10.11606 / issn. 2176 - 7793. v 25 i 4 p 203 - 265"]}

The Colombian weasel, Neogale felipei (Izor & de la Torre, 1978), is one of the most enigmatic and threatened carnivores in South America, with only six confirmed records in the Andes of Ecuador and Colombia. During a long-term trail camera survey conducted at Mesenia-Paramillo Natural Reserve, we recorded the northernmost occurrence of the species, which extends its distribution by approximately 120 km to the north from the nearest previously known locality in Colombia. We also provide some comments on its natural history.

Pimpla sumichrasti Cresson, 1874 (Fig. 1C) Pimpla sumichrasti Cresson, 1874: 400. Holotype ♀, Mexico (ANSP). Coccygomimus sumichrasti (Cresson) Townes, 1946: 44. Diagnosis. Head bright to rather dull yellow with black marks and mesosoma predominantly yellow or orange, sometimes with black marks; malar space narrow, less than 0.75�� as long as basal mandibular width, that of male less than 0.60�� basal mandibular width; mesoscutum yellow with three longitudinal black stripes; fore wing with an apical black spot and with Rs strongly sinuous; metasoma with laterotergites V narrow, less than 0.30�� as broad as long; TI of female rather slender, in profile evenly convex; female with tergites VI���VII almost entirely black. Biological note. Unknown (Gauld, 1991; Yu et al., 2012). Distribution. According to Yu et al. (2012): Argentina, Brazil, Costa Rica, Ecuador, Guatemala, Mexico, Paraguay, Peru and Venezuela. * Colombia., Published as part of P��dua, Diego G., Araujo, Rodrigo O. & Mazariegos, Luis A., 2019, Pimpla Fabricius (Hymenoptera: Ichneumonidae: Pimplinae) from Colombia, pp. 439-446 in Zootaxa 4683 (3) on page 441, DOI: 10.11646/zootaxa.4683.3.8, http://zenodo.org/record/3479049, {"references":["Cresson, E. T. (1874) Descriptions of Mexican Ichneumonidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 1873, 374 - 413.","Gauld, I. D. (1991) The Ichneumonidae of Costa Rica I. Memoirs of the American Entomological Institute, 47, 1 - 589.","Yu, D. S., van Achterberg, C. & Horstmann, K. (2012) World Ichneumonoidea 2011. Ottawa, Taxapad. Database on flash-drive. Available from: http: // www. taxapad. com (accessed 5 March 2018)"]}

Pimpla caerulea Brull��, 1846 (Fig. 1B) Pimpla caerulea Brull��, 1846. In Lepeletier: Histoire naturelle des insects Hym��nopt��res, 4: 101. des. Type: ♀, Brazil: ���Prov. De Rio Grande��� (MNHN). Coccygomimus caeruleatus Townes & Townes, 1946: 34. Coccygomimus caeruleatus caeruleus Townes & Townes, 1966: 24. Coccygomimus caeruleatus glaucus Townes & Townes, 1966: 25. Diagnosis. Clypeus weakly concave apically; mesosoma and metasoma metallic blue; wings blackish; fore wing with Rs more or less straight and cu-a opposite to Rs&M; metasoma with laterotergite V narrow, less than 0.30�� as long as wide; female with tergite I short and wide, strongly convex laterally; male with fore coxae white marked. Biological notes. Host is the Noctuidae moth Alabama argillacea (H��bner) (Porter, 1970). Distribution. According to Yu et al. (2012): Argentina, Bolivia, Brazil, Costa Rica, Ecuador, Guatemala, Mexico, Peru, Paraguay and Venezuela. * Colombia., Published as part of P��dua, Diego G., Araujo, Rodrigo O. & Mazariegos, Luis A., 2019, Pimpla Fabricius (Hymenoptera: Ichneumonidae: Pimplinae) from Colombia, pp. 439-446 in Zootaxa 4683 (3) on page 441, DOI: 10.11646/zootaxa.4683.3.8, http://zenodo.org/record/3479049, {"references":["Brulle, M. A. (1846) Tome Quatrieme. Des Hymenopteres. Les Ichneumonides. In: Lepeletier de Saint-Fargeau, A. (Ed.), Histoire Naturelles des Insectes, Paris, 1846, pp. 56 - 324.","Porter, C. C. (1970) A revision of the South American species of Coccygomimus (Hymenoptera, Ichneumonidae). Studia Entomologica, 13, 1 - 192.","Yu, D. S., van Achterberg, C. & Horstmann, K. (2012) World Ichneumonoidea 2011. Ottawa, Taxapad. Database on flash-drive. Available from: http: // www. taxapad. com (accessed 5 March 2018)"]}

Pimpla albomarginata Cameron, 1886 (Fig. 1A) Pimpla albo-marginata Cameron, 1886: 267. Holotype ♀, Mexico (BMNH). Coccygomimus albomarginatus (Cameron) Townes & Townes, 1966: 24. Diagnosis. Apex of clypeus deeply bilobed; malar space wide, longer than basal mandibular width, that males less than 0.60�� basal mandibular width; mesoscutum entirely black; postscutellum black; mesopleural suture weakly foveolate; propodeum with conspicuous posterolateral tubercles; fore wing Rs more or less straight and cu-a slightly distal to the base of Rs&M; coxae without black markings and fore coxa white markings; metasoma black and white banded with laterotergites V broad, more than 0.50�� as broad as long; tergite I of female short and broad, in profile strongly convex; tergite I in profile with moderately high blunt hump; sternite I with strongly produced swelling. Biological note. There is no host record for this species (Gauld, 1991; Yu et al., 2012). Distribution. According to Yu et al. (2012): Costa Rica, Mexico, Panama and Venezuela. * Colombia., Published as part of P��dua, Diego G., Araujo, Rodrigo O. & Mazariegos, Luis A., 2019, Pimpla Fabricius (Hymenoptera: Ichneumonidae: Pimplinae) from Colombia, pp. 439-446 in Zootaxa 4683 (3) on page 440, DOI: 10.11646/zootaxa.4683.3.8, http://zenodo.org/record/3479049, {"references":["Cameron, P. (1886) Hymenoptera. In: Godman, F. D. & Salvin, O. (Eds.), Biologia Centrali-Americana; or, Contributions to the knowledge of the fauna and flora of Mexico and Central America. Zoology, 1, pp. 241 - 328.","Gauld, I. D. (1991) The Ichneumonidae of Costa Rica I. Memoirs of the American Entomological Institute, 47, 1 - 589.","Yu, D. S., van Achterberg, C. & Horstmann, K. (2012) World Ichneumonoidea 2011. Ottawa, Taxapad. Database on flash-drive. Available from: http: // www. taxapad. com (accessed 5 March 2018)"]}

Andinobates cassidyhornae sp. nov. Dendrobates opisthomelas Silverstone 1975. Quebrada Arriba, bus stop 10 km by road from town of Andes, mountains near road (LACM 71962 - 70). Andinobates opisthomelas Brown & Twomey et al. 2011. pp 33 Plate 3, Figure O. Guatapé Antioquia, Colombia, because of a mistake in the final manuscript. The correct locality according to one of the authors of that paper (D.M.-V.), who actually took the photograph, is Carmen de Atrato, Chocó, Colombia. Holotype: An adult female (Figure 2) deposited in the amphibian collection of the Universidad de los Andes, Bogotá, Colombia, Andes-A 1095 (Field number LUMA 1001) is one of a series collected on March 30 th 2012 at the Mesenia-Paramillo Nature Reserve, Municipality of Andes, Department of Antioquia, Colombia, by R. Medina, G. Suárez & L. Mazariegos. Paratypes: Four adult females, and two adult males (Andes-A 1088–1091, and 1093–1095) collected by R. Medina, G. Suárez & L. Mazariegos. Locality data is the same as the type locality (see below). Type locality. Mesenia-Paramillo Nature Reserve, Vereda La Mesenia, about 12 km south of the municipality of Jardín but politically within the Municipality of Andes, (both in Departamento de Antioquia, Colombia), ca 5 ° 31´N, 75 ° 53´W at 2000 m elevation. Because of the heavy smuggling on dendrobatid frogs we refrain from providing more accurate coordinates. Etymology. This specific epithet cassidyhornae is a patronym in honor of Cassidy Horn, for her passionate interest in poison frogs and her generous contributions to the conservation of cloud forests in Colombia. Definition and diagnosis. A small-sized dendrobatid frog that we assign to the Andinobates bombetes species group (Brown & Twomey et al. 2011) based on the phylogenetic affinity (see Molecular and Phylogenetic Analyses) and on the following morphological characters: adult snout-vent length (SVL) et al. 2011). Andinobates cassidyhornae sp. nov. has an SVL of 19.03 ± 0.31 mm (mean±SD, N = 12 frogs), a bright red dorsum with the color extending onto the upper front and hind limbs; lower forearms and hind limbs are dark brown. Ventral coloration is black with bright red irregularly sized and spaced ovoid or ‘comma’ shaped blotches or spots. It can be externally distinguished from other species in the bombetes group by the distinctive color pattern: in A. cassidyhornae dorsum is bright red, and venter is black with well-defined bright red blotches or spots (Figure 4) vs. (1) in A. opisthomelas dorsum is red often with a posterior suffusion to brown, and venter is black with numerous white spots or reticulation (white-venter form) or venter is chocolate brown sometimes with red suffusion from the flanks (brown-venter form); (2) in A. virolinensis venter is whitish or bluish with black reticulation; (3) in A. bombetes the anterior half of dorsum exhibits bright red, yellow or rarely orange longitudinal and broad dorsolateral stripes; (4) in A. tolimensis the head is yellow, fading to brown towards the dorsum; (5) in A. dorisswansonae the dorsum is black or brown with red blotches, and the venter entirely black or black with few white or yellowish blotches; and (6) in A. daleswansoni the head is entirely red, and the body dull gold or brown. Measurements of the holotype (mm). The holotype measurements correspond to an adult female with a SVL of 18.99 mm; TL of 8.10; HaL of 4.58; HL of 4.85; HW of 6.29; GBW of 7.54; IOD of 2.31; HDT of 1.23; ED of 2.15; TSCN of 1.40; NED of 1.52; IND of 2.36; MTD of 0.75; W 3 FD of 0.78; W 3 F of 0.46; W 3 TD of 0.83; W 3 T of 0.47; W 4 TD of 0.87; and W 4 T of 0.49. The corresponding measurements of all specimens collected are shown in Table 2. Measurements Andes- Andes- Andes- Andes- Andes- Andes- Andes- Median SD A 1093 A 1088 A 1089 A 1090 A 1094 A 1091 A 1095 SVL 19.36 19.34 19.17 18.92 18.45 18.99 18.99 18.99 0.31 TL 8.59 8.52 8.23 8.51 8.50 8.66 8.10 8.51 0.20 HaL 4.59 4.67 4.67 4.60 5.14 4.91 4.58 4.67 0.21 HL 6.06 5.59 5.17 4.75 5.09 5.00 4.85 5.09 0.46 HW 5.38 5.99 5.94 5.75 6.20 5.90 6.29 5.94 0.30 GBW 8.50 8.85 8.13 7.11 7.56 7.20 7.54 7.56 0.66 IOD 2.38 2.30 2.12 1.85 2.26 2.14 2.31 2.26 0.18 HTD 1.41 1.17 0.98 0.96 1.05 0.97 1.23 1.05 0.17 ED 2.49 2.31 2.12 2.45 2.47 2.21 2.15 2.31 0.16 TSCN 1.78 1.34 1.26 1.22 1.54 1.35 1.40 1.35 0.19 NED 1.71 1.68 1.54 1.57 1.78 1.68 1.52 1.68 0.10 IND 2.47 2.33 2.32 2.42 2.84 2.56 2.36 2.42 0.18 MTD 0.84 0.72 0.67 0.60 0.89 0.91 0.75 0.75 0.12 W 3 FD 0.94 0.99 0.85 0.95 1.12 1.10 0.78 0.95 0.12 W 3 F 0.49 0.44 0.48 0.53 0.65 0.52 0.46 0.49 0.07 Description of the holotype: The head is slightly wider than longer, and is narrower than the body. Snout subovoid in dorsal view and truncated in lateral view. Canthus rostralis subovoid, loreal region flat and vertical. Nares situated much closer to the end of the snout than to the eyes, ovoid in shape and directed posterolaterally. Eyes large and prominent with a diameter of 11.3 % of SVL. The pupil is rounded and horizontally elliptical. Tympana and tympanic rings are in the posterolateral regions, ovoid and wider dorsoventrally, measuring 57.2 % of the diameter of the eyes. Supratympanic fold absent. Rounded choanae, not visible in ventral view, as they are completely covered by the maxillary arch. Vomerine, maxillary and premaxillary teeth are absent. Tongue is elongated, almost two times longer than wide; the posterior margin of the tongue is not indented and its posterior third is not adhered to floor of mouth. Hand relatively large (Figure 3), with a length equal to 24.1 % of the snout-vent length. The relative length of the fingers in increasing order of size is: IV The relative length of the toes, in increasing size order is I Coloration of holotype in life: Iris very dark brown; almost indistinguishable from black pupil. Nares encircled by black; margins of upper and lower jaws are very dark brown; tympana are black. Dorsum primarily bright scarlet red, sharply defined along the margins with a few irregularly scattered black speckles and small irregular black markings; in other specimens, black marks may reflect attacks by predators and should therefore not be considered an element of the dorsal coloration. Flanks are red laterally and black ventrolaterally with no gradient merging of the two colors as is seen in A. opisthomelas. Venter ground color black with irregular sharply contrasting bright scarlet red irregularly shaped blotches or spots. Upper forelimbs bright scarlet red and tinged brown where the upper and lower colorations meet, with lower forearm dark brown below the elbow; wrists and hands brown with tips of toes and fingers beige. Thigh mostly red, irregularly speckled with brown; shanks mostly brown tinged with red irregularly (Figure 2). Color in preservative (ethanol 70 %): The dark brown and black colors turn dull black to dark olive in preservative. Discs and tubercules on hands and feet, pupil and cornea become grey or nearly white with time. The predominantly bright scarlet red dorsal coloration and ventral red blotches and spots turn metallic olive; pattern remains clearly distinguishable in preservative (ethanol 70 %). Natural history: The natural history of this species is poorly known. We found individuals in areas covered with a thick layer of leaf litter and where abundant refuges were available. Males call regularly from the leaf litter or hidden amidst tree roots, throughout the day but prominently between 10 h– 14 h and after periods of rain. Most calling males were observed accompanied by a female. We also encountered several courting pairs during a visit in March and April 2012. Males carry 1–3 tadpoles (Figure 5) on their dorsum. Males of other species of the bombetes group are known to release their tadpoles in bromeliad water tanks. Some tadpoles of the new species were found in water within the inflorescence husks of Wettinia palms. Whether this species displays biparental care is unknown. Molecular and phylogenetic analyses. The final alignment consisted of 1119 bp (700 bp unambiguously aligned from the Cytb gene and 419 bp from the 16 S gene). The chosen partitioning strategy was as follows: 16 S unpartitioned under the GTR+ Γ model, and Cytb partitioned by codon position with K 80 +I, HKY+I and GTR+ Γ for the first, second, and third positions, respectively. Both ML and Bayesian phylogenies placed A. cassidyhornae sp. nov. as an independent, well-supported clade, separate from other species, within the bombetes species group. The two sequenced populations, the white- and the brown-venter forms, of A. opisthomelas formed a monophyletic albeit poorly supported group (Figure 6). The topology obtained is roughly consistent with the results of Brown & Twomey et al. (2011), but changes the relationships between A. virolinensis, A. tolimensis and A. bombetes, placing the first two as reciprocally monophyletic clades, and A. bombetes as the sister group of the two. However, the deeper phylogenetic relationships between the species in this group still require further study, since the obtained topology is highly polytomic and not well supported at this level. Within A. cassidyhornae sp. nov., individuals from the Mesenia-Paramillo Natural Reserve, Antioquia, cluster together in a monophyletic group, whereas individuals from other localities do not show evidence of genetic structuring. The SH tests rejected, at a very high level of statistical significance, two out of the three null topologies (H0) that placed A. cassidyhornae sp. nov. within a clade with A. opisthomelas (P A. virolinensis (P A. bombetes (P = 0.05134). Additionally, pairwise genetic distances (K 2 P) between A. cassidyhornae sp. nov. and other species within the bombetes species group ranged between 0.0320–0.0677, well within the range of interspecific distances (Figure 7) observed for the group (0.0216–0.1053), and about tenfold the intraspecific distances observed (0–0.0091). Altogether, the reconstructed phylogenetic tree, the topology (SH) tests, and pairwise genetic distances offer strong support for A. cassidyhornae sp. nov. being a distinct species within the bombetes species group. Bioacoustic analyses. All species’ calls within the Andinobates bombetes species group consist of long and atonal series of pulses sounding like a ‘buzz’ or rattle, which is often longer than 1 sec. The advertisement call of A. cassidyhornae sp. nov. (Figure 8) follows roughly the same pattern. Calls consist of 234.3 ± 20.3 (mean±SD) pulses, last 1.94 ± 0.26 s, and are often uttered as series of calls separated by regular silent intervals of 10.1 ± 2.1 s. The rise time is 50.0± 13.2 % of the call duration. The peak call frequency averages 4.32 ± 0.14 kHz and the frequency bandwidth 0.81 ± 0.40 kHz. Combining the calls of A. cassidyhornae sp. nov., A. opisthomelas (two localities), A. bombetes, and A. virolinensis, we found that call duration (linear regression, R 2 = 0.31, F= 13.9, P= 0.0008, N= 16 males), inter-call interval (R 2 = 0.17, F= 6.5, P= 0.0162), and the number of pulses per call (R 2 = 0.12, F= 4.3, P= 0.0472) decreased at higher temperatures. After removing the temperature effect by calculating regression residuals, the calls of A. cassidyhornae sp. nov. were clearly separated from the calls of the other species in a two-dimensional discriminant space (Figure 9, above, discriminant analysis, Wilks’ Lambda approxim. F= 13.9, PA. cassidyhornae sp. nov. from the call of any other species because the former were lower in peak frequency (F to enter= 39.6, standardized discriminant coefficient= 1.10, PA. cassidyhornae sp. nov. from the call of A. bombetes because the former were longer in duration (F to enter= 6.9, standardized discriminant coefficient= 0.97, P= 0.0013). Regarding pairwise species comparisons by univariate (Figure 9, below) tests, the advertisement call of A. cassidyhornae sp. nov. was lower in peak frequency than the call of A. opisthomelas (Tukey-Kramer HSD test, - 0.97 kHz, PA. virolinensis (- 0.71, PA. bombetes (- 0.53, PA. bombetes. Finally, its frequency bandwidth was wider compared to A. virolinensis (+ 0.40 kHz, P= 0.0220) and A. opisthomelas (+ 0.35 kHz, P= 0.0246). The strong among-species variation in call peak frequency could be partly attributed to concomitant variation in body size. Larger frogs usually produce calls at lower frequency values (e.g. Erdtmann & Amézquita 2009 for dendrobatid frogs) due to allometric constraints in larynx size. Indeed, increasing peak frequency is roughly related to decreasing body size in our study species: A. cassidyhornae sp. nov. is the largest species (mean±SD, 18.73 ± 0.22 mm, N = 5 recorded males, this study), followed by A. bombetes (17.76 ± 0.55, N = 28 males, Myers & Daly 1980), A. opisthomelas (16.80 ± 1.24, N = 26 males, Silverstone 1975), and A. virolinensis (16.72 ± 0.54, N = 127 males, Valderrama-Vernaza et al. 2009). We did not correct for body size effects on peak frequency, because we did not have all information on body size of recorded individuals. In any case, the difference in call frequency alone probably has important evolutionary implications. Across many frog species, ear sensitivity appears to match the peak frequency of the advertisement call (Capranica & Moffat 1983, see Amézquita et al. 2006, 2011 for examples on dendrobatid frogs). Thus, among-lineages differences in call frequency would imply a frequency mismatch between senders and receivers in the mate recognition signal, which could have promoted reproductive isolation between any pair of the Andinobates species we studied here. Distribution, habitat and ecology. At the type locality, the Mesenia-Paramillo Natural Reserve, Andinobates cassidyhornae sp. nov. was found in two fragments of heavily disturbed cloud forests. The approximately 1.52 hectare site is located on a steep montane slope with a gradient of 65 % (Figure 10). This mountain forest fragment is entirely surrounded by cattle grazing grasslands and agriculture crops. The forest fragment there has a dense, sometimes broken canopy with a complex stratification and emergent trees up to 20 m in height. The dominant canopy trees are Lauraceae (Nectandra acutifolia, Nectandra laurel, Aniba coto, Aiouea dubia, Aniba perutiles and Ocotea sp.), Oak (Quercus humboldtii) and Cedar (Cedrela montana). Shrubs and small trees in the families Ericaceae, Gesneriaceae, Melastomataceae, Piperaceae and Rubiaceae dominate the understory. Cyathea sp. tree ferns are common and epiphytes are dense on most trees dominated by the families Araceae, Bromeliaceae, Dryopteridaceae and Orchidaceae. Bromeliads are predominantly of the genus Guzmania. The forest floor is covered with abundant leaf litter and decomposing wood. Remains of inflorescences of Wettinia kalbreyerii palms are common, and serve as terrestrial water reservoirs for extended periods of time. Near daily (primarily evening and early morning) cloud cover of fog, mist and precipitation provide a very cool and humid mesic environment. No additional water sources, i.e. streams or springs were found there. The average annual rainfall is 2500 mm. The average annual regional temperature is 15.0° C. There are two conspicuous wet seasons beginning in March and lasting until May, and another extending from October through December. The species is also known from other three localities visited by one of the co-autors (D. M.-V., Figure 1) that are much less known than the type localitity. They, however, look roughly similar in topography and frogs’ microhabitat. Conservation status. Almost all the known localities for species in the A. bombetes group are within the 1200–2100 m elevational belt. In Colombia, the forests within this range have been severely degraded by intensive agriculture, remarkably coffee plantations, which is a first order national product. Andinobates cassidyhornae sp. nov. was found at four localities within 1800–2059 m elevation (Figure 1). The minimum area of the elevational range encompassing these localities, equivalent to the area achieved by the sum of the occupied grid squares (Figure 2 C in IUCN 2001), is between 200–300 km 2. However, to the best of or experience, the species distribution is sparsely patched including just few of the apparently suitable hills, probably occupying a minor fraction of the available habitat. In addition, most of the suitable forest below 2000 m elevation was cleared since many years ago at the type locality, which further limits the potential distribution of the new species. Unfortunately, most localities included in this study are exposed to severe degradation by intensive agriculture. Contamination of watersheds by pesticides, herbicides, and soi

Amézquita, Adolfo, Márquez, Roberto, Medina, Ricardo, Mejía-Vargas, Daniel, Kahn, Ted R., Suárez, Gustavo, Mazariegos, Luis (2013): A new species of Andean poison frog, Andinobates (Anura: Dendrobatidae), from the northwestern Andes of Colombia. Zootaxa 3620 (1): 163-178, DOI: http://dx.doi.org/10.11646/zootaxa.3620.1.8

Andinobates cassidyhornae sp. nov. Dendrobates opisthomelas Silverstone 1975. Quebrada Arriba, bus stop 10 km by road from town of Andes, mountains near road (LACM 71962 - 70). Andinobates opisthomelas Brown & Twomey et al. 2011. pp 33 Plate 3, Figure O. Guatap�� Antioquia, Colombia, because of a mistake in the final manuscript. The correct locality according to one of the authors of that paper (D.M.-V.), who actually took the photograph, is Carmen de Atrato, Choc��, Colombia. Holotype: An adult female (Figure 2) deposited in the amphibian collection of the Universidad de los Andes, Bogot��, Colombia, Andes-A 1095 (Field number LUMA 1001) is one of a series collected on March 30 th 2012 at the Mesenia-Paramillo Nature Reserve, Municipality of Andes, Department of Antioquia, Colombia, by R. Medina, G. Su��rez & L. Mazariegos. Paratypes: Four adult females, and two adult males (Andes-A 1088���1091, and 1093���1095) collected by R. Medina, G. Su��rez & L. Mazariegos. Locality data is the same as the type locality (see below). Type locality. Mesenia-Paramillo Nature Reserve, Vereda La Mesenia, about 12 km south of the municipality of Jard��n but politically within the Municipality of Andes, (both in Departamento de Antioquia, Colombia), ca 5 �� 31��N, 75 �� 53��W at 2000 m elevation. Because of the heavy smuggling on dendrobatid frogs we refrain from providing more accurate coordinates. Etymology. This specific epithet cassidyhornae is a patronym in honor of Cassidy Horn, for her passionate interest in poison frogs and her generous contributions to the conservation of cloud forests in Colombia. Definition and diagnosis. A small-sized dendrobatid frog that we assign to the Andinobates bombetes species group (Brown & Twomey et al. 2011) based on the phylogenetic affinity (see Molecular and Phylogenetic Analyses) and on the following morphological characters: adult snout-vent length (SVL) et al. 2011). Andinobates cassidyhornae sp. nov. has an SVL of 19.03 �� 0.31 mm (mean��SD, N = 12 frogs), a bright red dorsum with the color extending onto the upper front and hind limbs; lower forearms and hind limbs are dark brown. Ventral coloration is black with bright red irregularly sized and spaced ovoid or ���comma��� shaped blotches or spots. It can be externally distinguished from other species in the bombetes group by the distinctive color pattern: in A. cassidyhornae dorsum is bright red, and venter is black with well-defined bright red blotches or spots (Figure 4) vs. (1) in A. opisthomelas dorsum is red often with a posterior suffusion to brown, and venter is black with numerous white spots or reticulation (white-venter form) or venter is chocolate brown sometimes with red suffusion from the flanks (brown-venter form); (2) in A. virolinensis venter is whitish or bluish with black reticulation; (3) in A. bombetes the anterior half of dorsum exhibits bright red, yellow or rarely orange longitudinal and broad dorsolateral stripes; (4) in A. tolimensis the head is yellow, fading to brown towards the dorsum; (5) in A. dorisswansonae the dorsum is black or brown with red blotches, and the venter entirely black or black with few white or yellowish blotches; and (6) in A. daleswansoni the head is entirely red, and the body dull gold or brown. Measurements of the holotype (mm). The holotype measurements correspond to an adult female with a SVL of 18.99 mm; TL of 8.10; HaL of 4.58; HL of 4.85; HW of 6.29; GBW of 7.54; IOD of 2.31; HDT of 1.23; ED of 2.15; TSCN of 1.40; NED of 1.52; IND of 2.36; MTD of 0.75; W 3 FD of 0.78; W 3 F of 0.46; W 3 TD of 0.83; W 3 T of 0.47; W 4 TD of 0.87; and W 4 T of 0.49. The corresponding measurements of all specimens collected are shown in Table 2. Measurements Andes- Andes- Andes- Andes- Andes- Andes- Andes- Median SD A 1093 A 1088 A 1089 A 1090 A 1094 A 1091 A 1095 SVL 19.36 19.34 19.17 18.92 18.45 18.99 18.99 18.99 0.31 TL 8.59 8.52 8.23 8.51 8.50 8.66 8.10 8.51 0.20 HaL 4.59 4.67 4.67 4.60 5.14 4.91 4.58 4.67 0.21 HL 6.06 5.59 5.17 4.75 5.09 5.00 4.85 5.09 0.46 HW 5.38 5.99 5.94 5.75 6.20 5.90 6.29 5.94 0.30 GBW 8.50 8.85 8.13 7.11 7.56 7.20 7.54 7.56 0.66 IOD 2.38 2.30 2.12 1.85 2.26 2.14 2.31 2.26 0.18 HTD 1.41 1.17 0.98 0.96 1.05 0.97 1.23 1.05 0.17 ED 2.49 2.31 2.12 2.45 2.47 2.21 2.15 2.31 0.16 TSCN 1.78 1.34 1.26 1.22 1.54 1.35 1.40 1.35 0.19 NED 1.71 1.68 1.54 1.57 1.78 1.68 1.52 1.68 0.10 IND 2.47 2.33 2.32 2.42 2.84 2.56 2.36 2.42 0.18 MTD 0.84 0.72 0.67 0.60 0.89 0.91 0.75 0.75 0.12 W 3 FD 0.94 0.99 0.85 0.95 1.12 1.10 0.78 0.95 0.12 W 3 F 0.49 0.44 0.48 0.53 0.65 0.52 0.46 0.49 0.07 Description of the holotype: The head is slightly wider than longer, and is narrower than the body. Snout subovoid in dorsal view and truncated in lateral view. Canthus rostralis subovoid, loreal region flat and vertical. Nares situated much closer to the end of the snout than to the eyes, ovoid in shape and directed posterolaterally. Eyes large and prominent with a diameter of 11.3 % of SVL. The pupil is rounded and horizontally elliptical. Tympana and tympanic rings are in the posterolateral regions, ovoid and wider dorsoventrally, measuring 57.2 % of the diameter of the eyes. Supratympanic fold absent. Rounded choanae, not visible in ventral view, as they are completely covered by the maxillary arch. Vomerine, maxillary and premaxillary teeth are absent. Tongue is elongated, almost two times longer than wide; the posterior margin of the tongue is not indented and its posterior third is not adhered to floor of mouth. Hand relatively large (Figure 3), with a length equal to 24.1 % of the snout-vent length. The relative length of the fingers in increasing order of size is: IVColoration of holotype in life: Iris very dark brown; almost indistinguishable from black pupil. Nares encircled by black; margins of upper and lower jaws are very dark brown; tympana are black. Dorsum primarily bright scarlet red, sharply defined along the margins with a few irregularly scattered black speckles and small irregular black markings; in other specimens, black marks may reflect attacks by predators and should therefore not be considered an element of the dorsal coloration. Flanks are red laterally and black ventrolaterally with no gradient merging of the two colors as is seen in A. opisthomelas. Venter ground color black with irregular sharply contrasting bright scarlet red irregularly shaped blotches or spots. Upper forelimbs bright scarlet red and tinged brown where the upper and lower colorations meet, with lower forearm dark brown below the elbow; wrists and hands brown with tips of toes and fingers beige. Thigh mostly red, irregularly speckled with brown; shanks mostly brown tinged with red irregularly (Figure 2). Color in preservative (ethanol 70 %): The dark brown and black colors turn dull black to dark olive in preservative. Discs and tubercules on hands and feet, pupil and cornea become grey or nearly white with time. The predominantly bright scarlet red dorsal coloration and ventral red blotches and spots turn metallic olive; pattern remains clearly distinguishable in preservative (ethanol 70 %). Natural history: The natural history of this species is poorly known. We found individuals in areas covered with a thick layer of leaf litter and where abundant refuges were available. Males call regularly from the leaf litter or hidden amidst tree roots, throughout the day but prominently between 10 h��� 14 h and after periods of rain. Most calling males were observed accompanied by a female. We also encountered several courting pairs during a visit in March and April 2012. Males carry 1���3 tadpoles (Figure 5) on their dorsum. Males of other species of the bombetes group are known to release their tadpoles in bromeliad water tanks. Some tadpoles of the new species were found in water within the inflorescence husks of Wettinia palms. Whether this species displays biparental care is unknown. Molecular and phylogenetic analyses. The final alignment consisted of 1119 bp (700 bp unambiguously aligned from the Cytb gene and 419 bp from the 16 S gene). The chosen partitioning strategy was as follows: 16 S unpartitioned under the GTR+ �� model, and Cytb partitioned by codon position with K 80 +I, HKY+I and GTR+ �� for the first, second, and third positions, respectively. Both ML and Bayesian phylogenies placed A. cassidyhornae sp. nov. as an independent, well-supported clade, separate from other species, within the bombetes species group. The two sequenced populations, the white- and the brown-venter forms, of A. opisthomelas formed a monophyletic albeit poorly supported group (Figure 6). The topology obtained is roughly consistent with the results of Brown & Twomey et al. (2011), but changes the relationships between A. virolinensis, A. tolimensis and A. bombetes, placing the first two as reciprocally monophyletic clades, and A. bombetes as the sister group of the two. However, the deeper phylogenetic relationships between the species in this group still require further study, since the obtained topology is highly polytomic and not well supported at this level. Within A. cassidyhornae sp. nov., individuals from the Mesenia-Paramillo Natural Reserve, Antioquia, cluster together in a monophyletic group, whereas individuals from other localities do not show evidence of genetic structuring. The SH tests rejected, at a very high level of statistical significance, two out of the three null topologies (H0) that placed A. cassidyhornae sp. nov. within a clade with A. opisthomelas (P A. virolinensis (P A. bombetes (P = 0.05134). Additionally, pairwise genetic distances (K 2 P) between A. cassidyhornae sp. nov. and other species within the bombetes species group ranged between 0.0320���0.0677, well within the range of interspecific distances (Figure 7) observed for the group (0.0216���0.1053), and about tenfold the intraspecific distances observed (0���0.0091). Altogether, the reconstructed phylogenetic tree, the topology (SH) tests, and pairwise genetic distances offer strong support for A. cassidyhornae sp. nov. being a distinct species within the bombetes species group. Bioacoustic analyses. All species��� calls within the Andinobates bombetes species group consist of long and atonal series of pulses sounding like a ���buzz��� or rattle, which is often longer than 1 sec. The advertisement call of A. cassidyhornae sp. nov. (Figure 8) follows roughly the same pattern. Calls consist of 234.3 �� 20.3 (mean��SD) pulses, last 1.94 �� 0.26 s, and are often uttered as series of calls separated by regular silent intervals of 10.1 �� 2.1 s. The rise time is 50.0�� 13.2 % of the call duration. The peak call frequency averages 4.32 �� 0.14 kHz and the frequency bandwidth 0.81 �� 0.40 kHz. Combining the calls of A. cassidyhornae sp. nov., A. opisthomelas (two localities), A. bombetes, and A. virolinensis, we found that call duration (linear regression, R 2 = 0.31, F= 13.9, P= 0.0008, N= 16 males), inter-call interval (R 2 = 0.17, F= 6.5, P= 0.0162), and the number of pulses per call (R 2 = 0.12, F= 4.3, P= 0.0472) decreased at higher temperatures. After removing the temperature effect by calculating regression residuals, the calls of A. cassidyhornae sp. nov. were clearly separated from the calls of the other species in a two-dimensional discriminant space (Figure 9, above, discriminant analysis, Wilks��� Lambda approxim. F= 13.9, PA. cassidyhornae sp. nov. from the call of any other species because the former were lower in peak frequency (F to enter= 39.6, standardized discriminant coefficient= 1.10, PA. cassidyhornae sp. nov. from the call of A. bombetes because the former were longer in duration (F to enter= 6.9, standardized discriminant coefficient= 0.97, P= 0.0013). Regarding pairwise species comparisons by univariate (Figure 9, below) tests, the advertisement call of A. cassidyhornae sp. nov. was lower in peak frequency than the call of A. opisthomelas (Tukey-Kramer HSD test, - 0.97 kHz, PA. virolinensis (- 0.71, PA. bombetes (- 0.53, PA. bombetes. Finally, its frequency bandwidth was wider compared to A. virolinensis (+ 0.40 kHz, P= 0.0220) and A. opisthomelas (+ 0.35 kHz, P= 0.0246). The strong among-species variation in call peak frequency could be partly attributed to concomitant variation in body size. Larger frogs usually produce calls at lower frequency values (e.g. Erdtmann & Am��zquita 2009 for dendrobatid frogs) due to allometric constraints in larynx size. Indeed, increasing peak frequency is roughly related to decreasing body size in our study species: A. cassidyhornae sp. nov. is the largest species (mean��SD, 18.73 �� 0.22 mm, N = 5 recorded males, this study), followed by A. bombetes (17.76 �� 0.55, N = 28 males, Myers & Daly 1980), A. opisthomelas (16.80 �� 1.24, N = 26 males, Silverstone 1975), and A. virolinensis (16.72 �� 0.54, N = 127 males, Valderrama-Vernaza et al. 2009). We did not correct for body size effects on peak frequency, because we did not have all information on body size of recorded individuals. In any case, the difference in call frequency alone probably has important evolutionary implications. Across many frog species, ear sensitivity appears to match the peak frequency of the advertisement call (Capranica & Moffat 1983, see Am��zquita et al. 2006, 2011 for examples on dendrobatid frogs). Thus, among-lineages differences in call frequency would imply a frequency mismatch between senders and receivers in the mate recognition signal, which could have promoted reproductive isolation between any pair of the Andinobates species we studied here. Distribution, habitat and ecology. At the type locality, the Mesenia-Paramillo Natural Reserve, Andinobates cassidyhornae sp. nov. was found in two fragments of heavily disturbed cloud forests. The approximately 1.52 hectare site is located on a steep montane slope with a gradient of 65 % (Figure 10). This mountain forest fragment is entirely surrounded by cattle grazing grasslands and agriculture crops. The forest fragment there has a dense, sometimes broken canopy with a complex stratification and emergent trees up to 20 m in height. The dominant canopy trees are Lauraceae (Nectandra acutifolia, Nectandra laurel, Aniba coto, Aiouea dubia, Aniba perutiles and Ocotea sp.), Oak (Quercus humboldtii) and Cedar (Cedrela montana). Shrubs and small trees in the families Ericaceae, Gesneriaceae, Melastomataceae, Piperaceae and Rubiaceae dominate the understory. Cyathea sp. tree ferns are common and epiphytes are dense on most trees dominated by the families Araceae, Bromeliaceae, Dryopteridaceae and Orchidaceae. Bromeliads are predominantly of the genus Guzmania. The forest floor is covered with abundant leaf litter and decomposing wood. Remains of inflorescences of Wettinia kalbreyerii palms are common, and serve as terrestrial water reservoirs for extended periods of time. Near daily (primarily evening and early morning) cloud cover of fog, mist and precipitation provide a very cool and humid mesic environment. No additional water sources, i.e. streams or springs were found there. The average annual rainfall is 2500 mm. The average annual regional temperature is 15.0�� C. There are two conspicuous wet seasons beginning in March and lasting until May, and another extending from October through December. The species is also known from other three localities visited by one of the co-autors (D. M.-V., Figure 1) that are much less known than the type localitity. They, however, look roughly similar in topography and frogs��� microhabitat. Conservation status. Almost all the known localities for species in the A. bombetes group are within the 1200���2100 m elevational belt. In Colombia, the forests within this range have been severely degraded by intensive agriculture, remarkably coffee plantations, which is a first order national product. Andinobates cassidyhornae sp. nov. was found at four localities within 1800���2059 m elevation (Figure 1). The minimum area of the elevational range encompassing these localities, equivalent to the area achieved by the sum of the occupied grid squares (Figure 2 C in IUCN 2001), is between 200���300 km 2. However, to the best of or experience, the species distribution is sparsely patched including just few of the apparently suitable hills, probably occupying a minor fraction of the available habitat. In addition, most of the suitable forest below 2000 m elevation was cleared since many years ago at the type locality, which further limits the potential distribution of the new species. Unfortunately, most localities included in this study are exposed to severe degradation by intensive agriculture. Contamination of watersheds by pesticides, herbicides, and soi, Published as part of Am��zquita, Adolfo, M��rquez, Roberto, Medina, Ricardo, Mej��a-Vargas, Daniel, Kahn, Ted R., Su��rez, Gustavo & Mazariegos, Luis, 2013, A new species of Andean poison frog, Andinobates (Anura: Dendrobatidae), from the northwestern Andes of Colombia, pp. 163-178 in Zootaxa 3620 (1) on pages 168-177, DOI: 10.11646/zootaxa.3620.1.8, http://zenodo.org/record/216319

tesis que para obtener el grado de Doctorado en Ciencias Químicas, presenta Luis Rolando Mejia Mazariegos ; asesor J. Jesús Hernández Trujillo134 páginas : ilustracionesDoctorado en Ciencias Químicas UNAM, Facultad de Química, 2012