taxonID	type	description	language	source
4A5987B5315E1E36FAA0CB8DE51A842C.taxon	materials_examined	Type Material. Holotype male with labels: “ Rt. wing. [with tiny wing mounted on label] // Chatham I. / Galapagos. / VII- 1906 // Coll. by F. X. Williams // Neoryctes / galapagoensis / (G. R. Water.) ” and with my red holotype label. Type deposited at the U. S. National Museum (Washington, DC). Holotype. Male. Length 15.6 mm; width 8.9 mm. Color reddish brown. Head: Frons coarsely rugopunctate except for narrow, nearly smooth area mesad of each eye. Frontoclypeal suture weakly impressed, extending to each lateral margin. Clypeus subtriangular, apex narrowly subtruncate, surface transversely rugose; base either side of middle and just in front of frontoclypeal suture with small low, transverse tubercle. Mandibles large, exposed, arcuate up to apex where small, subapical “ tooth ” with longitudinal ridge beneath. Interocular width equals 4.8 transverse eye diameters. Antenna 9 - segmented, club subequal in length to antennomeres 2 – 6. Pronotum: Surface convex, lacking fovea or tubercles, punctures moderate in size and density, becoming slightly denser on apical third. Base with marginal bead except in front of scutellum where obsolete. Elytra: Surface with moderately large, moderately dense, shallow, glabrous punctures, some in indistinct rows. Apices bluntly rounded, appearing foreshortened, elytra together wider than long. Hind wings short, nearly vestigial, 4.0 mm in length. Pygidium: Surface rugulose in basal fifth and angles. Disc shiny, with a few small, glabrous punctures, becoming impunctate on apical third. In lateral view, surface weakly convex (nearly flat). Legs: Protibia tridentate. Metatibia at apex slightly expanded, subtruncate, with 15 short spinules. Venter: Prosternal process long, subrectangular, thick, slightly recurving, apex truncate. Parameres: Broadly rounded in basal half, distal half long, slender, and with pointed apices (Figs. 18 – 19).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315E1E36FAA0CB8DE51A842C.taxon	etymology	Etymology. The specific epithet is derived from the Greek anachoretes, meaning a hermit or recluse. It is used here as a masculine noun in apposition to re-emphasize, along with the generic name, the reclusiveness of this rare species.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315E1E36FAA0CB8DE51A842C.taxon	distribution	Distribution. Adelaeus anachoretus is known only from San Cristóbal Island in the Galápagos Archipelago, Ecuador. San Cristóbal was the first island in the archipelago visited by Charles Darwin in 1835 while on the Beagle expedition. Locality Record. ECUADOR (1): GALÁPAGOS (1): San Cristóbal Island (Fig. 20). Temporal Distribution. July (1).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315E1E36FAA0CB8DE51A842C.taxon	diagnosis	Diagnosis. The characters listed in the generic diagnosis, greatly reduced flight wings, the unique form of the parameres, and the occurrence in the Galápagos Islands will serve to distinguish this species. The only other New World dynastine with this form of parameres is Tomarus rostratus Dupuis. Natural History. The holotype of A. anachoretus lacks detailed collecting information. It may have been collected during the 1905 – 1906 California Academy of Sciences expedition to the Galápagos Islands and then possibly exchanged or gifted to the USNM. The July 1906 material collected by F. X. Williams is mentioned in Van Dyke’ s (1953) overview of the Coleoptera of the Galápagos Islands, but San Cristóbal Island is not mentioned as among the localities for the 18 dynastine specimens collected by Williams. Since no other specimens have been found in the intervening 108 years, it could be surmised that A. anachoretus is an extremely rare endemic that has not been observed in previous collecting efforts due to a secretive life style, or that it is now extinct due possibly to the introduction of predatory invasive vertebrates. Cook et al. (1995) observed that the then-known four Neoryctes species in the Galápagos Islands were all rare, due possibly to the inaccessibility of the high elevation habitat, short period of adult nocturnal activity that coincided with rain, or predation by introduced rats, pigs, and ants. The volcanic islands of the Galápagos Archipelago are approximately 800 – 1,000 km west of mainland Ecuador. All of the islands are volcanic in origin and semiarid with the lowlands having a short wet season, while the highlands catch fog and rain clouds and are moister (Van Dyke 1953). Tye et al. (2002) observed that isolation of the islands has led to a high degree of endemism for vertebrates, and Peck (2005) suggested that limitations for establishment of colonizing beetles is more related to the strongly seasonal, semiarid climate of the large, lowland areas of the islands and the prevalence of large areas of young and unweathered lava substrates that are not conducive to beetles. Peck (2005) recorded the beetle fauna of the Galápagos Islands at 56 families, 297 genera, and 486 species with 266 endemic species, 110 indigenous species, and 110 introduced species. Previously, Neoryctes was the only endemic genus of scarabs in the Galápagos Islands. Peck (2005) concluded that generic endemism and species diversification is strongly correlated with secondary loss of flight wings after colonization, and that most colonization has not been followed by much speciation. Both A. anachoretus and N. linelli inhabit San Cristóbal (Fig. 20), and both have reduced hind wings.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315D1E30FB7CCE4FE78C818C.taxon	type_taxon	Type Species. Adelaeus anachoretus Ratcliffe, here designated.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315D1E30FB7CCE4FE78C818C.taxon	description	Description. Body form short, suboval (Fig. 17), elytra together wider than long. Length about 16 mm. Color reddish brown. Clypeus subtriangular, apex narrowly subtruncate. Interocular distance equals about 5 transverse eye diameters. Antenna with 9 antennomeres, club subequal in length to antennomeres 2 – 6. Mandibles exposed, rounded, with small, subapical tooth. Pronotum with punctures moderate in density and size; base margined except in front of scutellum where bead obsolete. Elytra with moderately dense, shallow punctures, striae indistinct. Hind wings reduced. Pygidium nearly flat, rugose across base, disc with few small punctures. Protibia tridentate. Metatibia at apex subtruncate. Prosternal process thick, subrectangular, slightly recurved, apex truncate.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315D1E30FB7CCE4FE78C818C.taxon	distribution	Distribution. San Cristóbal (Chatham) Island, Galápagos Islands, Ecuador.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315D1E30FB7CCE4FE78C818C.taxon	diagnosis	Diagnosis. The only other Dynastinae occurring on the Galápagos Islands are five species of the endemic genus Neoryctes, and one might naturally expect for this new species to belong to that genus. Adelaeus anachoretus, while superficially similar to Neoryctes species in body form, is unique and differs from Neoryctes as follows: clypeal apex narrowly subtruncate (narrowly bidentate in Neoryctes); mandibles rounded with small, subapical tooth (mandibles with external edge rounded, entire in Neoryctes); prosternal process long, subrectangular, thick, slightly recurved, apex truncate (process short, laminate, subtriangular in Neoryctes); propygidium glabrous and lacking stridulatory bands (propygidium finely setose and with two longitudinal stridulatory bands in Neoryctes); pygidium in males nearly flat (strongly convex in males of Neoryctes); and parameres long, slender, and with acute apices (Figs. 18 – 19) (parameres generally subrectangular in caudal view in Neoryctes) (Cook et al. 1995). Only N. linelli occurs sympatrically on San Cristóbal (Chatham) Island, but that beetle is much larger, has quadridentate protibiae, and the parameres are broadly subrectangular. Neoryctes species and A. anachoretus have hind wings so greatly reduced that they cannot be used for flight, and it is possible that the short, suboval body form seen in both is a result of reduced wings. In size and gestalt, A. anachoretus also resembles in body form species of Eremobothynus Ohaus. The three species of Eremobothynus occur in Argentina and have a central frontoclypeal tubercle or horn, a declivous pronotum in males, functional flight wings, and generally subtriangular parameres, none of which are character states shared with A. anachoretus.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315D1E30FB7CCE4FE78C818C.taxon	etymology	Etymology. The generic epithet is derived from the Greek stem adelos, meaning unknown or obscure and is used here as a masculine noun to mean “ that which is unseen ” in reference to the long hidden nature of this genus.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531551E38FB4CCE1AE6278111.taxon	description	663 although 130 species have been described since this work. Consequently, the keys in Endrödi’ s manual are now incomplete because there exist 44 % more species than are in the keys. Cyclocephala species occur from extreme southeastern Canada south to Argentina and Chile and in the West Indies. One species has been introduced into Australia. Most of the species occur in Central and South America. Species in the genus may be recognized by a clypeus that is subparallel at its base and with the sides converging anteriorly to a rounded, parabolic, subtruncate, or emarginate apex; antenna with 8 – 10 segments and the club longer in the males of some species; and protarsus in the males distinctly enlarged with the median (inner) claw much larger than the outer claw. The protarsus in females is simple and not enlarged. Adult Cyclocephala species are nocturnal, and they are attracted to lights at night. Only a few larvae have been described, and those seem to feed on the roots of grasses. Adults of some tropical species are known to feed upon and pollinate inside the flowers of aroids and certain palms.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531561E3EF8CFCB62E6798627.taxon	materials_examined	Type Material. H o l o t y p e m a l e l a b e l e d “ BOLIVIA: Santa Cruz / Flora & Fauna Lodge / 3.7 km SSE Buena Vista / 17 ° 29 ′ 55 S, 63 ° 39 ′ 9 W / 17 – 19 November 2006 / B. Ratcliffe and M. Jameson ” and with my red holotype label. Allotype female labeled “ BOLIVIA: Santa Cruz Dist. / 4 km SSE Buena Vista Hotel / Flora & Fauna, 17 - 20. X. 2011 / 17 ° 29 ′ S, 63 ° 39 ′ W, 300 – 400 m / Ex: b / l; Wappes and Skillman ” and with my red allotype label. One male paratype with same data as allotype. One paratype male labeled “ BOLIVIA / Santa Cruz / Feb. 1956 / G. Pinckert ” and with my yellow paratype label. One paratype male labeled “ BOLIVIA: Santa Cruz, 3.7 km / SSE Buena Vista, Hotel Flora / and Fauna 405 m., 5 - 15 - XI- 2001 / 17 ° 29.949 ′ S; 63 ° 33.152 ′ W / M. C. Thomas & B. K. Dozier / tropical transition forest ” and with my yellow paratype label. One paratype female labeled “ BOLIVIA / Santa Cruz Dept. / October 2011 ” and with my yellow paratype label. One paratype female labeled “ BOLIVIA: Santa Cruz Dept. / Potrerillos del Guenda, Snake / Farm, 21 - 24. X. 2011; 370 – 400 m / 17 ° 40 ′ S 63 ° 27 ′ W; Ex blacklight / Coll. J. E. Wappes & F. Skillman ” and with my yellow paratype label. Holotype and allotype deposited at the University of Nebraska State Museum (Lincoln, NE). Paratypes deposited as follows: one to the U. S. National Museum (Washington, DC), one to the Florida State Collection of Arthropods (Gainesville, FL), one to the Fabien Dupuis Collection (Saint- Chamond, France), and two to the Brett C. Ratcliffe Collection (Lincoln, NE). Holotype. Male. Length 15.3 mm; width 7.6 mm. Color completely testaceous except for black apices of protibial teeth, femora, and tibiae. Head: Frons and clypeus with small, dense punctures. Clypeus with apex subtruncate, narrowly reflexed. Interocular width equals 3.0 transverse eye diameters. Antenna with 10 segments, club slightly longer than antennomeres 2 – 7. Pronotum: Surface with small, moderately dense punctures, punctures becoming slightly denser on sides. Base lacking marginal bead. Elytra: Surface with punctate striae and with randomly arranged punctures in intervals, especially first; punctures moderately large, glabrous. Pygidium: Surface completely, densely scabriculous, glabrous. In lateral view, surface weakly, regularly convex. Legs: Protibia tridentate, teeth equidistant from one another. Protarsus enlarged; tarsomere 5 large, curved, with strong, longitudinal carina on median surface; median claw large, strongly bent at base, apex narrowly cleft. Metatarsus distinctly longer than metatibia. Venter: Prosternal process long, columnar, apex with fringe of long setae and obliquely flattened into transversely suboval disc with anterior half elevated into a raised, transverse “ button. ” Parameres: Sides subparallel, constricted well before the apex, apices evenly rounded (Figs. 2 – 3). Allotype. Female. Length 14.0 mm; width 7.2 mm. As holotype except in the following respects: Elytra: Epipleuron (ventral view) of females gradually thickened to and then abruptly constricted at level of abdominal sternite 1; in dorsal view, lateral margin of elytra with large swelling at about middle. Pygidium: Surface completely rugulopunctate, glabrous, nearly flat in lateral view. Legs: Metatarsus subequal in length to metatibia.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531561E3EF8CFCB62E6798627.taxon	description	Variation. Males (3 paratypes). Length 13.6 – 14.1 mm; width 6.8 – 7.1 mm. The paratypes do not differ significantly from the holotype. One specimen has the antennal club subequal in length to antennomeres 2 – 7. One specimen has short, sparse setae along the base of the pygidium. Females (2 paratypes). Length 13.5 – 13.9 mm; width 6.9 – 7.4 mm. The paratypes do not differ significantly from the allotype.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531561E3EF8CFCB62E6798627.taxon	etymology	Etymology. I am pleased to name this species after my French colleague, Fabien Dupuis, in recognition of his numerous contributions to the taxonomy of Dynastinae.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531561E3EF8CFCB62E6798627.taxon	distribution	Distribution. To date, C. dupuisi is known only from two locations in Santa Cruz Department, Bolivia, but, no doubt, it occurs elsewhere in the department where there is suitable forest habitat. Locality Records. BOLIVIA: SANTA CRUZ: Flora and Fauna Lodge (Fig. 4), the Snake Farm. Temporal Distribution. February (1), November (6).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531561E3EF8CFCB62E6798627.taxon	diagnosis	Diagnosis. The parameres of C. dupuisi closely resemble those of Cyclocephala occipitalis Fairmaire from southern Brazil and Argentina. However, C. dupuisi has a testaceous frons and vertex (black or piceous in C. occipitalis) and an antennal club in the male that is subequal to or only slightly longer than antennomeres 2 – 7 (as long as or longer than the entire stem in C. occipitalis). In addition, C. dupuisi is a larger beetle at 14 mm plus, whereas C. occipitalis is 13 mm or smaller. Cyclocephala dupuisi males will key as far as couplet 435 / 436 in Endrödi (1985), where the choices for Cyclocephala flavipennis Arrow and Cyclocephala sanguinicollis Burmeister do not conform to the characters of C. dupuisi, especially the sculpturing of the pygidium and shape of the parameres. Females will key to couplet 357 in Endrödi’ s (1985) female key, where, again, the character states are different: lateral margin of elytra swollen at about middle in C. dupuisi and swollen behind the middle in C. flavipennis; pygidium completely rugulopunctate in C. dupuisi and sparsely punctate in C. flavipennis. But remember that there are now at least 130 species that have been described since Endrödi’ s work that are not in his key. Natural History. The holotype was collected at a light trap in secondary forest near the Hotel Flora and Fauna (Fig. 4). The other specimens were probably taken at lights as well.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531581E2BFAD4CE65E6EF8647.taxon	materials_examined	Type Material. Holotype male with handwritten label: “ La Esperranza / Cundinamarca / Dpt. 24) Parameres, lateral view. Colombia / SA. 1250 m. Oct. ′ 35 / RPRoba # EH ” and with my red holotype label. Holotype deposited at the University of Nebraska State Museum (Lincoln, NE). Holotype. Male. Length 20.3 mm; width 8.7 mm. Color black. Head: Surface smooth except for small, dense punctures just mesad of each eye and on each ocular canthus. Vertex with deep, smooth pit. Frons with long (2.2 mm), erect, slightly recurving horns; in frontal view, horns becoming slightly broader on apical half. Clypeus subtriangular, apex acute and strongly reflexed; carina extending from clypeal apex to base of each horn. Interocular width equals 4.6 transverse eye diameters. Antenna 10 - segmented, club distinctly longer than antennomeres 2 – 7. Pronotum: Surface with large, deep, moderately dense punctures; punctures small and sparse immediately either side of median furrow. Median longitudinal furrow deep, broad, becoming wider at anterior end, extending from near base to apical declivity; surface of furrow with dense, large, often confluent punctures. Base with complete marginal bead. Elytra: Surface with 6 punctate striae between suture and humerus; punctures large, ocellate, separated by about 1 puncture diameter or less between large punctures or about 2 diameters between smaller punctures. Intervals distinctly convex, smooth, all subequal in height. Sides with 5 rows of similar punctures. Pygidium: Surface densely punctate; punctures moderately large, setigerous; setae tawny, moderately dense, most moderate in length. In lateral view, surface convex. Legs: Protibia quadridentate, teeth subequally spaced. Apex of metatibia with spine-like upper tooth and with 3 short, thick spinules in crenulations below. Venter: Prosternal process short, thick, triangularly columnar, apex bluntly rounded, posterior surface with distinct prominence at base. Last sternite with sparse, moderately large punctures along basal margin, elsewhere sparsely micropunctate. Parameres: Simple, suboval, constricted at about middle, apices rounded (Figs. 23 – 24).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531581E2BFAD4CE65E6EF8647.taxon	etymology	Etymology. The specific epithet is derived from the Latin cornutus, meaning “ bearing horns or horned. ” Hence, “ the Hemiphileurus with horns ” is in reference to the proportionately larger horns (for a Hemiphileurus species) of the holotype.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531581E2BFAD4CE65E6EF8647.taxon	distribution	Distribution. Hemiphileurus cornutus is known only from the type locality just to the west of Bogotá in the department of Cundinamarca in Colombia. Locality Record. COLOMBIA (1): CUNDINA- MARCA: La Esperanza (Fig. 25). Temporal Distribution. October (1).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531581E2BFAD4CE65E6EF8647.taxon	diagnosis	Diagnosis. Hemiphileurus cornutus is characterized by the surface of the head nearly smooth, with distinct horns in the major male holotype; pronotum with a long and deep median furrow; elytral intervals distinctly convex and all of equal height; prosternal process small, thick, and subtriangularly columnar; protibiae quadridentate; and form of the parameres. Compared to the other 13 Hemiphileurus species from Colombia, H. cornutus is the only one with quadridentate protibiae and so is readily distinguished ... until an additional new species is discovered with similar protibiae. Hemiphileurus cornutus will key as far as couplet 45 / 46 in Endrödi (1985), where the choices for H. microps (Burmeister) and H. illatus do not conform to the characters of H. cornutus. However, there are now 29 species that have been described since Endrödi’ s work that are not in his key. Natural History. Nothing is known of the immature stages or life history of this species. The specimen was collected at an elevation of 1,250 m.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531581E36F89ECE65E0F9842C.taxon	description	In Colombia, there are now 14 species of Hemiphileurus with the description of this new species: H. blandini Dupuis, H. caliensis Endrödi, H. cornutus Ratcliffe, new species, H. curvicornis Dupuis and Dechambre, H. depressus (F.), H. dejeani (Bates), H. deslislesi Ratcliffe, H. elbitae Neita and Ratcliffe, H. gibbosus Dupuis and Dechambre, H. hiekei Chalumeau, H. laticollis (Burmeister), H. rugulosus Endrödi, H. variolosus (Burmeister), and H. vicarius Prell. The genus continues to grow in number of species as new localities are explored and sampled and as unidentified specimens residing in research collections are studied. Compare Endrödi’ s (1985) listing of 24 known species at that time with the 51 species now recognized. The genus is characterized by having species whose adults have pointed mandibles simply curved externally, clypeus triangularly pointed, frons with tubercles (usually) or horns (occasionally) arising far from the lateral margin of the head, pronotum usually with a longitudinal furrow but lacking an anterior fovea or strong declivity, and apex of the metatibia with a single, large tooth on the upper angle. Most of the species are moderate in size and range from 16 – 24 mm in length. Our knowledge of the life history and habits of these beetles remains virtually non-existent. This is a result of the secretive habits of most of the adults and larvae. Adults may be attracted to lights at night, and this is where virtually all specimens in collections were taken. But until our search pattern for collecting goes beyond light traps (i. e., extensive excavation of rotting logs), many Hemiphileurus species will remain rare in collections. The larval stages for only three species have been described: H. illatus (LeConte) (Ritcher 1966), H. dispar Kolbe (Ocampo and Morón 2004), and H. elbitae (Neita and Ratcliffe 2010). Larvae live in decaying wood, where they presumably feed on the wood and associated fungi.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E3CFB7AC8EAE5C2876F.taxon	description	The single species in the genus is characterized by a reddish brown color; frons with a single tubercle; clypeus strongly contracted towards apex; mandibles exposed and evenly rounded; antenna 10 - segmented with short club; pronotum simply convex; long prosternal process; quadridentate protibia; and metatibia at apex with spinules. The biology of H. tuberculatus is unknown. the Brazilian goddess of the wilderness who protects animals from unethical hunters.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E3CFB7AC8EAE5C2876F.taxon	distribution	Distribution. Stenocrates caiporae is known only from the village of Estirão do Equador that is situated on the Rio Yavari on the border of Brazil’ s Amazonas state with Peru. Locality Record. BRAZIL: AMAZONAS: Estirão do Equador (Fig. 8). Temporal Distribution. November (2).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E3CFB7AC8EAE5C2876F.taxon	diagnosis	Diagnosis. The form of the parameres is a necessary distinguishing character in Stenocrates species since their external body morphology is usually so similar. Accordingly, S. caiporae is not here incorporated into Endrödi’ s (1985) key. The parameres of S. caiporae are remarkably similar to those of Stenocrates haackae Ratcliffe (Figs. 9 – 10), which was described from central Amazonian Brazil. But there the similarities end. Stenocrates caiporae and S. haackae are compared in Table 1. Natural History. As is the case with most other species of Stenocrates, nothing is known of the life history of S. caiporae.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E32FADECA46E6168127.taxon	materials_examined	Type Material. Holotype female with handwritten label: “ PERU: Cusco / Salineras de Maras / S 13 ° 18 ′ 08 ″ W 72 ° 9 ′ 15 ″ / 5 Jan 2012, 3000 m / M. Jameson and L. Russell // Found dead in / salt catchment ” and with my red holotype label. Type deposited at the University of Nebraska State Museum (Lincoln, NE). Holotype. Female. Length 30.0 mm; width 14.5 mm. Color dark reddish brown. Head: Frons with large, dense, deep, confluent punctures. Frontoclypeal region at center with short, transversely arcuate tubercle. Clypeus triangular, apex narrowly rounded, not reflexed, surface transversely rugose. Mandibles exposed, large, evenly rounded. Interocular width equals 4.0 transverse eye diameters. Antenna 10 - segmented, club subequal in length to antennomeres 2 – 7. Pronotum: Surface convex, lacking fovea or tubercles, weakly crazed, anterior fifth half with moderately large, dense, deep punctures. Remainder of surface with moderately large punctures sparse to moderate in density. Base with thick marginal bead. Elytra: Sutural stria a row of deep, elongate punctures. Surface completely, densely punctate, punctures mostly large, glabrous. Pygidium: Disc shiny, with small, sparse, glabrous punctures. Base with narrow, transverse band of, setigerous punctures; punctures small to moderate, and setae long, reddish brown. In lateral view, surface convex. Legs: Protibia quadridentate. Metatibia at apex expanded, truncate and minutely crenulate with 17 stout spinules. Venter: Prosternal process moderately long, laminate, apex subtruncate.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E32FADECA46E6168127.taxon	etymology	Etymology. The specific epithet is derived from the Greek words halos for salt and philia for fondness. It is used here in reference to the salt catchments in which it was found. This is not to imply that this species actually has a predilection for salty habitats, but that it was simply found dead in such a place.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E32FADECA46E6168127.taxon	distribution	Distribution. Hiekeianus halophilus is known only from the area of the Maras salt pan area in Cusco Department, Peru. Locality Record. PERU: CUSCO: Salineras de Maras (Fig. 12). Temporal Distribution. January (1).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531521E32FADECA46E6168127.taxon	diagnosis	Diagnosis. The female of H. halophilus is described as new because it is so different from the females of H. tuberculatus. The differences between H. halophilus and H. tuberculatus are summarized in Table 2. In size and gestalt, H. halophilus also resembles Barutus hartmanni Ratcliffe from Panama and Costa Rica, but in B. hartmanni the clypeal apex is broader, the mandibles are weakly lobed, the frontoclypeal region has a binodose tubercle, the mentum is lanceolate rather than constricted at the subapex, the last abdominal sternite is subtruncate, and the prosternal process is large, thick, and rounded as opposed to the laminate, subtruncate process in H. halophilus. Natural History. The single holotype was found dead, and so nothing is known of its life history.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315C1E32F977CB63E72A8131.taxon	description	All species in the genus are characterized by a length of 13 – 27 mm; short and broad body form; reddish brown color; clypeus narrowed to a bidentate apex; mandibles exposed and evenly rounded; antenna 9 - or 10 - segmented with a short club; pronotum simply convex; prosternal process small and subtriangular; hindwings reduced; propygidium with two longitudinal stridulatory ridges; pygidium convex, especially in males; protibia tridentate or quadridentate; and parameres generally subrectangular with broadly lanceolate apices. All the species are rare, and their life history is unknown.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315C1E33FABACB4EE0058433.taxon	materials_examined	Type Material. Holotype male labeled: “ Cawley Mt. / Alt. Vii- 06 / Albermarle I / Galapagos Is // Coll. by / F. X. Williams ” and with my red holotype label. Right wing mounted on small card beneath specimen. Allotype female with same data and wing mounted beneath specimen. Types deposited at the California Academy of Sciences (San Francisco, CA). Holotype. Male. Length 16.0 mm; width 8.7 mm. Color reddish brown. Head: Frons completely, coarsely rugopunctate, punctures large. Frontoclypeal ridge distinctly elevated, arcuate in anterior view. Clypeus subtriangular, apex weakly bidentate; surface rugopunctate, punctures moderate in size. Ocular canthus lacking acute lateral process but with small, distinct lobe at base. Mandibles large, exposed, arcuate on external edge. Interocular width equals 4.5 transverse eye diameters. Antenna 9 - segmented, club subequal in length to antennomeres 2 – 6. Pronotum: Surface convex, lacking fovea or tubercles, punctures moderately large, moderate in density, becoming slightly sparser on basal third. Base with complete marginal bead. Elytra: Surface with punctures moderate in size and density, shallow, glabrous, some in distinct rows. Apices bluntly rounded, appearing foreshortened, elytra together as wide as long. Hind wings 8.7 mm long. Pygidium: Surface with minute, sparse punctures, shiny. In lateral view, surface strongly convex. Legs: Protibia quadridentate, basal tooth nearly obsolete. Metatibia at apex slightly expanded, subtruncate, with about 20 short spinules (some absent). Venter: Prosternal process short, triangular, laminate. long, subrectangular, thick, slightly recurving, apex truncate. Parameres: Together subrectangular, apices broadly expanded, median edge of each paramere just below middle with prominent tooth (Figs. 14 – 15). Allotype. Female. Length 16.8 mm; width 8.8 mm. As holotype except in the following respects: Pronotum: Punctures slightly denser. Hind wing 8.3 mm in length. Pygidium: Convex but not strongly so.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315C1E33FABACB4EE0058433.taxon	etymology	Etymology. The specific epithet is Latin that indicates “ similar to ... ”, in reference to the similarity between N. similis from Isla Isabela and Neoryctes linelli (Mutchler) from Isla San Cristóbal.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315C1E33FABACB4EE0058433.taxon	distribution	Distribution. Neoryctes similis is known only from Isabela Island (formerly Albermarle) in the Galápagos Archipelago, Ecuador (Fig. 16). Locality Records. ECUADOR (2): GALÁPAGOS (2): Isla Isabela. Temporal Distribution. July (2).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B5315C1E33FABACB4EE0058433.taxon	diagnosis	Diagnosis. This species will key out to couplet 3 in Cook et al. (1995) because of the length of the wing (8.7 mm), but then only parts of each of the remaining two choices conform to the specimen. Neoryctes moreti Dechambre has reduced eyes and an acute lateral process on the ocular canthus, whereas N. similis has neither; N. moreti has a reduced medial process on the parameres, whereas this process is prominent in N. similis. In the second part of couplet 3, Neoryctes williamsi Cook has the hind wing greatly reduced and lacking an anal fold, whereas N. similis has a larger wing with an anal fold. The parameres of N. similis are most similar to those of N. linelli, but in that species the wings are longer than 10 mm. Neoryctes species have variable character states (Cook et al. 1995), and so identifications can be problematical. Cook et al. (1995) was aware of, but did not see, the Cawley Mountain specimens referred to in Van Dyke (1953), and the new species described here is undoubtedly those specimens. They were in the collections of the California Academy of Sciences and placed in a series with an old header label of N. galapagoensis (Waterhouse) (L. Saylor determination). Henry Howden arranged for the loan from the Academy’ s Neoryctes specimens for the revision by Cook et al. (1995), but it seems that not all were sent since the Cawley Mountain material was not included in the revision. Natural History. Nothing is known of the life history for any of the five Neoryctes species.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531501E3EF897CC7CE7188045.taxon	description	Unlike most other species in the tribe Cyclocephalini, the males do not have enlarged protarsal claws. Species in the genus are also characterized by a relatively short, subtrapezoidal clypeus that has its apex truncate to slightly emarginate; a distinct frontoclypeal suture; three pairs of punctate striae on the elytra; and the meso- and metafemora strongly flattened. The antenna is 10 - segmented with a small club. Most of the species are externally similar to one another and difficult to tell apart, and so great reliance is made on the form of the male parameres for identification. This genus and Hemiphileurus Kolbe are possibly the most difficult genera of Dynastinae in the Americas with which to work. Most females not associated with males cannot usually be identified with reliability. Because of the high number of cryptic species in Stenocrates, new species continue to be discovered and described. The immature stages remain unknown for all Stenocrates species. Life history information is also lacking. The adults are attracted to lights. Endrödi (1966, 1985) provided the last synopsis of the genus, but 11 new species have been described since that time. These remain unincorporated in any key, and so a new generic synopsis and key is needed.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531501E3FFAA9CA5EE0D58159.taxon	materials_examined	Type Material. Holotype and single male paratype with handwritten label: “ Estirão do Equador / Rio Yavari – Amazo. / XI – 1960 – J. Hidasi ” and with my red holotype and yellow paratype labels, respectively. Holotype deposited at the University of Nebraska State Museum (Lincoln, NE, USA). Paratype deposited in the B. C. Ratcliffe collection (Lincoln, NE, USA). Holotype. Male. Length 25.7 mm; width 11.8 mm. Color dark reddish brown. Head: Frons shiny, with small, sparse punctures in basal half, punctures becoming slightly larger mesad of each eye and near frontoclypeal suture. Frontoclypeal suture distinctly impressed, ridge in front of suture vaguely carinulate either side of middle and obsolete at middle. Clypeus transversely rugose, apex shallowly emarginate, apical rim thickened. Interocular width equals 2.2 transverse eye diameters. Antenna with 10 segments, club subequal in length to antennomeres 2 – 7. Pronotum: Surface shiny, with sparse micropunctures, punctures becoming slightly larger along lateral margins. Lateral margins with thick marginal bead, base without marginal bead. Elytra: Surface shiny, with punctate sutural stria and 2 pairs of distinct discal striae and 1 pair of striae behind humerus; each stria comprised of large, deep, closely adjacent punctures. First broad interval with an irregular, almost double row of large, sparse punctures. Pygidium: Surface shiny, completely and densely punctate, punctures moderately large, glabrous. In lateral view, surface regularly convex. Legs: Protibia tridentate, teeth subequally spaced. Metatarsus shorter than metatibia. Venter: Prosternal process long, thick; in lateral view, columnar, apex broadly rounded; in ventral view, apex suboval. Parameres: Widest at about middle, with a subapical “ tooth ” on each side, and apices rounded (Figs. 6 – 7).	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531501E3FFAA9CA5EE0D58159.taxon	description	Variation. Male (1 paratype). Length 25.0 mm; width across humeri 12.0 mm. The paratype does not differ significantly from the holotype. Head: Interocular width equals 2.5 transverse eye diam- eters. Pygidium: Apical third becoming slightly less densely punctate. The female remains unknown.	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
4A5987B531501E3FFAA9CA5EE0D58159.taxon	etymology	Etymology. The specific epithet is derived from Caipora, who, in the Tupi-Guarani mythology, is	en	Ratcliffe, Brett C. (2014): A New Genus and Species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other New Species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a Revised Key to the Genera of New World Pentodontini. The Coleopterists Bulletin 68 (4): 663-680, DOI: 10.1649/0010-065X-68.4.663, URL: http://dx.doi.org/10.1649/0010-065x-68.4.663
