identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
4E574931E2725135B5A689A4E7DD70E0.text	4E574931E2725135B5A689A4E7DD70E0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onthophagus humboldti	<div><p>Onthophagus humboldti sp. nov. Figures 1, 2a, c, e, 3a, c, 5, 6, 7, 9</p><p>Type locality.</p><p>Costa Rica. Prov. Puntarenas. Buenos Aires, P.N. La Amistad. Tres Colinas.</p><p>Type deposition.</p><p>Museo Nacional de Costa Rica, Santo Domingo de Heredia, Costa Rica.</p><p>Type material.</p><p>Holotype male, pinned, with genitalia in a separate microvial. Original label: "Costa Rica. Provincia Puntarenas. Buenos Aires, Parque Nacional La Amistad. Tres Colinas. 2100-2200 m. 27-29 Febrero 2008. A. Solís, M. Moraga. Trampa Foso. L S 343850 565700." "HOLOTYPE/ Onthophagus humboldti Kohlmann, Solís, Alvarado [red printed label]".</p><p>Other material.</p><p>Paratypes. (8 males, 4 females). "Costa Rica. Provincia Puntarenas. Buenos Aires, Parque Nacional La Amistad. Tres Colinas. 2100-2200 m. 27-29 Febrero 2008. A. Solís, M. Moraga. Trampa Foso. L S 343850 565700."</p><p>Diagnosis.</p><p>Elytra as long as or shorter than pronotum (Fig. 1), due to brachyptery (Fig. 3c). Broad clypeal horn bifurcation (Fig. 2a); pygidium and apex of elytra with evident setae; clypeal margin indented at junction with clypeo-genal suture (Fig. 2e).</p><p>Description.</p><p>Holotype. Male (Fig. 1), length 7.2 mm; maximum width 4.3 mm. Oval, shining reddish black. Centre of the clypeus projected forming a slender bifid horn (Figs 1, 2c); genae projected forming an angle (Fig. 2e), genal sutures almost effaced; head punctures coarse, regular, dense, becoming finer and sparser towards the center; clypeal carina absent, vertex carina substituted by two vertical asymmetric platelets, modestly developed, and obliquely oriented; eyes two times longer than wide and separated by eight times the eye width.</p><p>Pronotum (Fig. 2c) very convex, lateral margins with a small and irregular fovea, not lineal; lateral pronotal margins bordered by a deep sulcus, anterior and basal borders margined; pronotal surface reticulate and covered by dense, regular, coarse, annular, and deep punctures without setae; pronotal projection well-developed (Figs 1, 2c), forming a broad bilobed plate slightly bent downwards, with a depressed area antero-centrally, and having clear antero-lateral margins; anterior angles projected as long, slender, and curved projections (Figs 1, 2c); pronotal base with a sulcus extending forward one third its length; scutellum not visible between the base of the elytra.</p><p>Elytra convex, with clear margins and without a humeral callus; with eight well-marked striae, fine and clearly impressed and with crenulating punctures; intervals clearly punctured, punctures big and dense, not aligned, bearing short, stiff setae along the lateral and apical margins; microsculpture reticulate and regular. Wing brachypterous, measuring 0.75 mm (Fig. 3c). Pygidium moderately shiny and shagreen, margined border, with big, coarse, annular punctures bearing short and stiff setae. Aedeagus as Fig. 3a.</p><p>Mesosternum with evident annular punctures bearing no setae. Metasternum shagreen and finely punctured, more coarsely laterally, basal third with a sulcus. Abdominal segments shagreen and finely punctured.</p><p>Fore femur long, slender, and punctured; meso- and metafemur short and elongate, light yellow. Fore tibia long, slender and arched (Fig. 1); with four external teeth; tibial spur elongated, straight, pointed, deflexed anteriorly, extending to second tarsal segment. Middle- and hind femur light yellow at middle.</p><p>Female, length 6.3 mm; maximum width 3.6 mm. It is similar to the male and varies in having a clypeus not forming a horn, clypeus shagreen, genae not projected as teeth, with a head frons keel, two small platelet projections at head vertex, no pronotal projection, no projected pronotal anterior angles, fore tibia short, fore femur short, last abdominal sternite broad.</p><p>Variation.</p><p>Length 5.6 to 7.2 mm. Width 3.2 to 4.3 mm. Small males do not have the bifid clypeal horn, just a small erect lamella; vertex platelets forming a small projection; anterior pronotal angles not projected, pronotal projection forming a small carina. Body color varying from black to piceous red.</p><p>Etymology.</p><p>This species is dedicated in honor of Friedrich Wilhelm Heinrich Alexander von Humboldt, Prussian geographer, explorer, and naturalist, commemorating the 250th anniversary of his birth. He is widely recognized for fathering the work on physical and plant geography, which laid the foundation for the development of modern biogeography.</p><p>Taxonomic considerations.</p><p>Kohlmann and Solís (2001) report the existence of 39 species of Onthophagus for Costa Rica. This new species would increase their numbers to 40. Onthophagus humboldti sp. nov. belongs to the Onthophagus dicranius Bates species group, as defined by Kohlmann and Solís (2001).</p><p>Onthophagus humboldti sp. nov. will key out to O. micropterus Zunino &amp; Halffter, 1981, in Kohlmann and Solís´ key (2001). It can be easily differentiated by the following characteristics: In males clypeal horn slender at middle and very bifurcated at apex (Fig. 2a) ( O. humboldti sp. nov.) versus broad at middle and notched at apex (Fig. 2b) ( O. micropterus); genae projected forming an angle (Fig. 2e) ( O. humboldti sp. nov.) versus genae projected forming a tooth (Fig. 2f) ( O. micropterus); vertex platelets forming a carina ( O. humboldti sp. nov.) versus a pointed projection ( O. micropterus); anterior lateral angles of pronotum projected as long, slender, and curved projections (Fig. 2c) ( O. humboldti sp. nov.) versus a short, curved projection (Fig. 2d) ( O. micropterus); pronotal central forward projection well-developed, forming a broad bilobed plate slightly bent downwards (Fig. 2c) ( O. humboldti sp. nov.) versus a bilobed plate projecting forward (Fig. 2d) ( O. micropterus). In females: vertex platelets forming a carina ( O. humboldti sp. nov.) versus a pointed projection ( O. micropterus).</p><p>Geographical distribution.</p><p>This species is so far only known from the area of Tres Colinas, near Buenos Aires, in the province of Puntarenas (Fig. 5). It has been collected from 2100 to 2200 m altitude in the month of February in lower montane rain forest.</p><p>Chorological affinities.</p><p>Onthophagus humboldti sp. nov. is endemic to the Cordillera de Talamanca and is the tenth known brachypterous Onthophagus species to be described worldwide. A closely related species, O. micropterus, is also distributed in the Cordillera de Talamanca (Fig. 6), from 2100 to 3000 m altitude in tropical mountain rainforest and has been collected from October to February.</p><p>Biogeography.</p><p>This species belongs to the O. dicranius species group, as established by Kohlmann and Solís (2001). This group of species has extra-American affinities, in which Howden and Gill (1993) indicate that the American fauna of Onthophagus is the result of invasive species from East Asia and that the O. dicranius group presents characters in common with New Guinea species. This agrees with the hypothesis originally proposed by Zunino and Halffter (1988), which points out for the supraspecific groups of American Onthophagus, an origin of its lineages, which in the case of the current representatives is distributed in East or Southeast Asia; and for this case, the Asian representation of the ancestral line, like the American one, has its distribution present in the humid tropics. On the other hand, the O. dicranius species group has its present-day center of diversity in tropical North America and relatives in South America (Zunino and Halffter 1997; Kohlmann and Solís 2001).</p><p>This situation seems to be in congruence with the boreotropical distribution hypothesis (Wang 1961; Wolfe 1975; Lavin and Luckow 1993; Xiang and Soltis 2001; Davis et al. 2002), where current flora groups show a tropical disjunct distribution, generally centered in America, Africa, and tropical Asia. This hypothesis is based on the observation of the existence of tropical broadleaf forests during the Early Paleogene (in old Stratigraphy terminology, Early Tertiary) at high latitudes in regions that are currently temperate, directed by a Late Paleocene-Early Eocene thermal maxima (ca. 52 ma, Zachos et al. 2001) and that many current angiosperm temperate taxa have evergreen relatives in subtropical rainforests (Axelrod 1966). This proposal then suggests the existence of northern bridges that were once at lower latitudes, such as the Bering Bridge during the Early Paleogene and the North Atlantic Bridge during the Eocene, which may have served as migration routes for groups of organisms that currently present intercontinental disjunct distributions. This hypothesis suggests that a taxon with a present-day center of diversity in tropical North America, and with an early Paleogene fossil record from any region there, has a high probability of having sister-group relatives in the Paleotropics and derived relatives in South America (Lavin and Luckow 1993).</p><p>This pattern of distribution would clarify those proposed by Halffter (Halffter and Morrone 2017) for the "Mexican Transition Zone" in particular one of them, the so-called "Paleoamerican Dispersion Pattern" (Halffter 1964). This pattern of dispersion corresponds to northern taxa that arrived in North America from Eurasia, and has been subdivided by Halffter et al. (1995) into four variants, where one of them, called the "Paleoamerican Tropical Pattern", corresponds to species found in the lowlands of the tropics and at medium altitudes, their distribution being very similar to that of the Neotropical pattern, but their affinities are with the Old World taxa. Halffter et al. (1995, 2008) placed the Onthophagus clypeatus and&gt; Onthophagus dicranius species groups of the genus Onthophagus within this pattern.</p><p>Actually, the groups of species mentioned above are congruent with the typical characteristics of the so-called boreotropical distribution. Therefore, the aforementioned distribution variant, the "Paleoamerican Tropical Pattern", seems to be the same with the boreotropical distribution and it is proposed here to use the term boreotropical distribution from now on as it is a more complete and well-founded concept, besides being an older one. This pattern has been studied and characterized at very fine phylogenetic and biogeographic analysis levels in animal and plants (Lidgard and Crane 1990; Xiang and Soltis 2001; Davis et al. 2002; Feng et al. 2009; Guo et al. 2012; Ye et al. 2016).</p></div>	https://treatment.plazi.org/id/4E574931E2725135B5A689A4E7DD70E0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Kohlmann, Bert;Solis, Angel;Alvarado, Guillermo E.	Kohlmann, Bert, Solis, Angel, Alvarado, Guillermo E. (2019): Description of Onthophagus humboldti and Uroxys bonplandi, two new scarab beetles (Coleoptera, Scarabaeidae, Scarabaeinae) from Costa Rica, with notes on tropical mountain brachyptery and endemicity. ZooKeys 881: 23-51, DOI: http://dx.doi.org/10.3897/zookeys.881.38026, URL: http://dx.doi.org/10.3897/zookeys.881.38026
CD2B9D8B870A5AA787E67D6936B3E89F.text	CD2B9D8B870A5AA787E67D6936B3E89F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Uroxys bonplandi	<div><p>Uroxys bonplandi sp. nov. Figures 2h, 3b, 4, 5</p><p>Type locality.</p><p>Costa Rica. Guanacaste. Sector Santa María, path to the cone of the Santa María, part of the Rincón de la Vieja volcanic massif, 1565 m.</p><p>Type deposition.</p><p>Museo Nacional de Costa Rica, Santo Domingo de Heredia, Costa Rica.</p><p>Type material.</p><p>Holotype male, pinned, with genitalia in a separate microvial. Original label: "Costa Rica. Provincia Guanacaste. Sector Santa María, Sendero a Pico Volcán Santa María . 1565 m. 2 Diciembre 2017. Col. Sergio Salas Ríos . Biocol. 10.8039N, 85.3281W." "HOLOTYPE/ Uroxys bonplandi Kohlmann, Solís, Alvarado [red printed label]".</p><p>Other material.</p><p>Paratypes (18 males, 25 females). "Costa Rica. Provincia Guanacaste. Sector Santa María, Sendero a Pico Volcán Santa María . 1565 m. 2 Diciembre 2017. Col. Sergio Salas Ríos . Biocol. 10.8039N, 85.3281W (6 males, 10 females). " Tilarán Bosque Nuboso Santa Elena. 1600 m. 26 Noviembre - 8 Diciembre 1999. J. Rodríguez Trampa de Luz. L N 258000 45000" (1 female). " Provincia Puntarenas. Monteverde Zona Protectora Arenal-Monteverde. Parcela Brillantes. 1500-1600 m. 17-19 Junio 2009. A. Solís, J.D. Gutiérres . Trampa Foso. L N 252009 450981" (4 males, 2 females), “13– 1600 m. 10°18'N, 84°48'W. Univ. California EAP 1991" (1 female). "Est. La Casona. 1520 m. Reserva Biológica Monteverde. N. Obando. Octubre 1991. L N 253250 449700" (2 males, 2 females), "Septiembre 1990 (1 male), 29 Nov - 17 Diciembre 1994, K. Martínez, L N 253200 449700" (2 males, 1 female). " Provincia Alajuela. San Ramón . Zona Protectora Arenal-Monteverde. Parcela El Valle. 1600-1700 m. 16-18 Jun 2009. A. Solís, J.D. Gutiérrez . Trampa Foso. L N 255970 452538" (3 males, 9 females).</p><p>Diagnosis.</p><p>Anterior of frons evenly convex, without carina or groove, with a dimple or transversely rugose; clypeal margin indented at junction with clypeogenal suture; dorsal ocular area twice as long as wide, distance between eyes five times eye width; pronotum evenly convex, sides angled near middle; elytral apex of the second to fourth intervals forming an oblique keel (Fig. 2h); basal sulcus of pygidium sinuate; fore tibial spur slender and deflexed distally.</p><p>Description.</p><p>Holotype. Male, length 7.4 mm; maximum width 3.8 mm. Elongate oval, shining reddish black (Fig. 4). Clypeus bidentate, slightly indented immediately laterad of teeth; teeth broadly triangular and strongly reflexed (Fig. 4). Head surface with a small dimple at the center, distinct small punctures throughout. Clypeogenal suture distinct; clypeal margin distinctly indented at intersection of suture (Fig. 4); genal margins broadly rounded (Fig. 4). Frons weakly convex, with very slight, broad indentations. Dorsal ocular areas approximately twice as long as wide at posterior edge of canthus (12 to 14 facets wide at that point), distance between ocular areas approximately five times their width.</p><p>Pronotum at median angulation as wide as elytra; lateral edges of pronotum produced into prominent angles (Fig. 4), strongly sinuate on lateral view, posterior two-thirds of margin nearly vertical; pronotum weakly convex medially; surface densely covered with fine, deep punctures; median longitudinal sulcus feebly indicated in posterior third; lateral fovea in form of crenulated longitudinal deep grove three-fourths length of pronotum (Fig. 4), not extending to either anterior or posterior margin, with cluster of coarse punctures in posterior third; pronotum margined basally, with adjacent row of large longitudinal punctures (Fig. 4).</p><p>Elytron moderately convex, clearly punctate (faintly in Uroxys dybasi Howden &amp; Young, 1981), humeral umbone small; striae distinct but shallow, with distinct punctures evenly spaced for most of length of each stria, seventh stria extending three-fifths length of elytron; posterior tenth of first stria furrowed; intervals flat, slightly flattened and constricted, not produced, except at the apex of the second to fourth intervals forming an oblique keel (Fig. 2h) (sharp straight keel in the third interval in dybasi, Fig. 2g).</p><p>Meso- and metasternum clearly punctate (faintly in dybasi); meso-metasternal suture medially moderately angulate anteriorly, moderately angulate laterally, three times farther from anterior margin of mesosternum than from mesocoxal cavity; metasternum swollen, with distinct median posterior depression.</p><p>Ventral abdominal segments two to five of equal length medially, each only slightly shorter medially than sixth; sixth slightly longer laterally than medially; anterior margins with small punctures (big crenulated punctures in dybasi). Pygidium strongly convex, faintly punctate, twice as wide as long; sulcus surrounding disc deep basally, shallow elsewhere; margin formed of same width apically and laterally; sulcus basally very slightly arcuate toward apex on each side of midline.</p><p>Fore tibia elongate with inner margin broadly curved (Fig. 4); outer margin with three teeth in apical third, teeth approximately equidistant, basal tooth somewhat reduced and more broadly triangular (Fig. 4); apex of fore tibia with short, narrow, rounded, deflexed projection at inner corner, projection approximately half length of tibial spur. Tibial spur elongated, straight, pointed, extending to fourth tarsal segment. Fore femur gradually tapering distally; middle femur with a faint ventral posterior triangular projection at apical third (evident projection in dybasi); hind femur with a well-developed ventral posterior swelling at apical third; posterior margin of hind trochanter continuous with posterior margin of femur.</p><p>Female, length 6.9 mm; maximum width 3.6 mm. It is similar to the male and varies in having a rugose clypeus, lateral edges of pronotum produced into less prominent angles. Elytral apex without oblique keels. Fore femur and fore tibia not as long. Middle and hind femur without a projection or swelling at apical third.</p><p>Variation .</p><p>Length 5.7 to 7.6 mm. Width 3.2 to 4.1 mm. The center of the head might have a small dimple and/or also a slight transverse rugosity.</p><p>Etymology.</p><p>This species is dedicated in honor of Aimé Jacques Alexandre Goujaud Bonpland, French naturalist, physician, and botanist, member of the scientific expedition that accompanied Humboldt to Spanish America.</p><p>Taxonomic considerations.</p><p>Solís and Kohlmann (2013) report the existence of 12 species of Uroxys for Costa Rica. This new species would increase their numbers to 13. Due to its great similarity, we here propose that Uroxys bonplandi sp. nov. represents the sister species of U. dybasi Howden &amp; Young, 1981.</p><p>Uroxys bonplandi sp. nov. will key out to U. dybasi in Solís and Kohlmann´s (2013) key. It can be easily differentiated by the following characteristics: Uroxys bonplandi sp. nov. is consistently bigger (5.7 to 7.6 mm) than its sister species (4.3 to 5.6 mm), U. dybasi . It can also be separated by the clear punctures in thorax and elytra in bonplandi sp. nov. (faint in dybasi). In males: elytral apex of the second to fourth intervals forming an oblique keel in bonplandi sp. nov. (Fig. 2h) (sharp straight keel in the third interval in dybasi, Fig. 2g), meso- and metasternum clearly punctate on bonplandi sp. nov. (faintly in dybasi), anterior margins of ventral abdominal segments with small punctures in bonplandi sp. nov. (big crenulated punctures in dybasi), and middle femur with a faint ventral posterior triangular projection at apical third in bonplandi sp. nov. (evident projection in dybasi).</p><p>Geographical distribution.</p><p>Uroxys bonplandi sp. nov. has been collected so far in the Cordillera de Guanacaste and the Cordillera de Tilarán (Fig. 5). It is a mountain species distributed from 1520 to 2200 m of altitude and has been collected in the following life-zones: wet tropical forest (premontane transition), lower montane rain forest, lower montane wet forest, premontane rainforest, and premontane wet forest. It has been collected from June to February.</p><p>Chorological affinities.</p><p>Uroxys bonplandi sp. nov. coincides with U. dybasi in being distributed along the Guanacaste and Tilarán mountain ranges. (Fig. 5) This last species has been also reported from mountain forests from Panama in the Cordillera de Chiriquí and in Costa Rica in the Cordillera Central and Talamanca (Fig. 5), being distributed between 600 and 1700 m and collected throughout the whole year. U. bonplandi sp. nov. represents also the first known endemic species of Uroxys for Costa Rica.</p><p>Another related species is Uroxys tacanensis Delgado &amp; Kohlmann, 2007, known only from its type locality, the Tacaná volcano, at the border of Mexico and Guatemala, living in cloud forest at 2000 m altitude (Delgado and Kohlmann 2007). No other species of this group has been yet collected in the intermediate areas. They are all montane species.</p></div>	https://treatment.plazi.org/id/CD2B9D8B870A5AA787E67D6936B3E89F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Kohlmann, Bert;Solis, Angel;Alvarado, Guillermo E.	Kohlmann, Bert, Solis, Angel, Alvarado, Guillermo E. (2019): Description of Onthophagus humboldti and Uroxys bonplandi, two new scarab beetles (Coleoptera, Scarabaeidae, Scarabaeinae) from Costa Rica, with notes on tropical mountain brachyptery and endemicity. ZooKeys 881: 23-51, DOI: http://dx.doi.org/10.3897/zookeys.881.38026, URL: http://dx.doi.org/10.3897/zookeys.881.38026
