Revision of the Genus Prionacalus (Coleoptera: Cerambycidae: Prioninae: Prionini)

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1 Revision of the Genus Prionacalus (Coleoptera: Cerambycidae: Prioninae: Prionini) Author(s): Antonio Santos-Silva Ziro Komiya and Eugenio H. Nearns Source: The Coleopterists Bulletin, 67(3): Published By: The Coleopterists Society URL: BioOne ( is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne s Terms of Use, available at terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.

2 The Coleopterists Bulletin, 67(3): REVISION OF THE GENUS PRIONACALUS WHITE, 1845 (COLEOPTERA: CERAMBYCIDAE: PRIONINAE: PRIONINI) ANTONIO SANTOS-SILVA Museu de Zoologia, Universidade de São Paulo Caixa Postal , São Paulo, São Paulo, BRAZIL ZIRO KOMIYA , Shimouma, Setagaya-ku Tokyo, , JAPAN AND EUGENIO H. NEARNS USDA, Department of Entomology, Smithsonian Institution National Museum of Natural History Washington, DC , U.S.A. ABSTRACT Prionacalus White, 1845, a genus restricted to western South America from Colombia to northern Argentina, is revised. Of the 15 species currently described, seven are considered as distinct and redescribed herein. The following five new synonymies are proposed: Prionacalus gunteri Gahan, 1894 is removed from synonymy with Prionacalus buckleyi Waterhouse, 1872 and proposed as a new synonym of Prionacalus cacicus (White, 1845); Prionacalus simonsi Waterhouse, 1900 is a new synonym of Prionacalus atys White, 1850; Prionacalus giovannii Hüdepohl, 1985 is a new synonym of Prionacalus whymperi Bates, 1892; Prionacalus trigonodes Bates, 1892, currently a synonym of P. buckleyi, is proposed as a new synonym of P. whymperi; and P. buckleyi is a new synonym of Prionacalus iphis White, The following earlier synonymies are accepted: Prionacalus emmae Kolbe, 1902 and Prionacalus whitei Waterhouse, 1900 are synonyms of P. buckleyi (and consequently, synonyms of P. iphis); syntype female of Psalidognathus wallisi Taschenberg, 1870 is a synonym of P. cacicus. Waterhouse (1872) is considered the author of the designation of the lectotype for Prionacalus iphis. The designation of a lectotype for P. whymperi by Quentin and Villiers is considered invalid. Lectotypes are designated for P. whymperi, P. buckleyi, and Prionacalus uniformis Waterhouse, A key to species is provided and all species are figured. Key Words: lectotype, longhorn beetles, South America, synonymy, taxonomy Prionacalus White, 1845 currently includes nine species (Bezark and Monné 2013) distributed in Ecuador, Peru, Colombia, and Argentina. Members of this genus are medium to large size, flightless prionines of the tribe Prionini, found primarily in the Andes Mountains, or around them, often at altitudes above 2,000 m. Notwithstanding Prionacalus is a small genus, the identification of its species is particularly difficult, mainly because they show considerable variability. Nearly all species are also uncommon in the natural history museums, making the study of the genus problematic. The specimens deposited in these institutions frequently were collected many years ago and often do not have detailed data label. On the other hand, specimens are somewhat common in private collections, probably due to thier high commercial value. Relatively common mistakes in the identification of the species make this study necessary to clarify the situation of each species. This revision is based on original type material of nearly all species described until now. MATERIAL AND METHODS The collection acronyms used in this study are as follows: BMNH The Natural History Museum, London, UK DDPC Diethard Dauber Private Collection, Austria 201

3 202 THE COLEOPTERISTS BULLETIN 67(3), 2013 DHPC EMEC IRSN MLUH MPCO MZSP NMPC SNSD SMNS USNM ZKCO ZMHB ZMUC ZSMC Daniel Heffern Private Collection, Houston, TX, USA Essig Museum of Entomology, University of California, Berkeley, CA, USA Institute Royal des Sciences Naturelles de Belgique, Brussels, Belgium Martin-Luther-Universität, Wissenschaftsbereich Zoologie, Halle, Germany Giuseppe and Valentino Marazzi Private Collection, Italy Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil National Museum (Natural History), Prague, Czech Republic Senckenberg Naturhistorische Sammlungen Dresden, Dresden, Germany Staatliches Museum für Naturkunde, Stuttgart, Germany National Museum of Natural History, Washington, DC, USA Ziro Komiya Collection, Tokyo, Japan Museum für Naturkunde, Berlin, Germany Zoological Museum, University of Copenhagen, Copenhagen, Denmark Zoologische Staatssammlung des Bayerischen Staates, Munich, Germany As numerous translations from Latin, German, and French are included in this work, we have preceded each section with (translation) and placed the translated text in quotes to clarify where it begins and ends. The species name in the bibliography under each species follows the original spelling. The citation listed after the country/state in Geographical Distribution refers to the first record of the species for that place. TAXONOMIC HISTORY White (1845) described the genus Prionacalus as a subgenus of Prionus Geoffroy, 1762, to allocate his new species Prionus (Prionacalus) cacicus, based on three male specimens. According to White (1845), those specimens were from Mexico: In a small collection of insects from Mexico, purchased last year by Mr. Gray for the British Museum from M. Hartweg, there are three specimens of a subgenus of Prionidae, at first sight with very considerable resemblance to the Psalidognathus modestus, Fries, Vetensk. Akad. Handl. 1833, p t. 9. f. 3, agreeing with that species in many particulars, but to me appearing distinct. White (1845) provided figures of two male specimens (pl. VIII: fig. 1) and one female specimen (pl. VIII: fig. 2) of P. cacicus. In the same work, White suggested that Psalidognathus modestus Fries, 1833 also belonged in the genus Prionacalus. However, P. modestus is a true Psalidognathus Gray, 1831, as seen by the length of the head (excluding mandible) and prothorax distinctly shorter than elytral length in males (similar in Prionacalus). Blanchard (1845) divided the subfamily Prioninae Latreille, 1802 into several groups including Psalidognathites with a single genus: Psalidognathus. Prionacalus was not mentioned by Blanchard (1845), and it is unclear if that work appeared before the publication of White (1845). According to Bousquet et al. (2009), the name Psalidognathites is unavailable (ICZN 1999: Art ). However, according to Bouchard et al. (2011): Psalidognathites Blanchard, 1845b: 138 [stem: Psalidognath-]. Type genus: Psalidognathus Gray, Comment: original vernacular name unavailable (Art ): subsequently used in Latinized form but not generally attributed to Blanchard (1845b). Reiche (1850a), in the meeting of the Académie des Sciences ofparisheldon9january1850, affirmed that Prionacalus is a synonym of Psalidognathus (translation): Mr. Reiche communicated a note about the genus Psalidognathus that, according to him, currently consists of four species, the P. Friendii Gray, Anim. kingd (superbus Fries, Vetensk. Akad. Handl., 1850), modestus Fries, id., erythrocerus Reiche, Rev. zool., 1840, all three from Colombia, and the P. cacicus White, An. and Magaz. of nat. history, 1845, from Mexico. Mr. Reiche dealt more specifically of P. cacicus, which Mr. White considered the type of a new genus, that of Prionocalus [sic], but that he [Reiche] believes must be retained in the genus Psalidognathus itself, and on which he gave details of interesting habits. According to Reiche (1850b) (translation): Mr. White, naturalist attached to the British Museum, had the first knowledge of this species [P. cacicus]. Comparing it with Psal. Friendii, the only species that existed at the Museum, he noticed the differences that he thought sufficient to establish a new genus named Prinocalus [sic]. Those differences consist mainly in the greater brevity of the abdomen and elytra; the fusion of the latter and absence of wings. By induction and from the description in Vetensk. Akad. Handl. 1833, he thought that it was his duty to bring to this new genus Psalid. modestus of this author [Fries]. This species is well known to the entomologists of Paris, and none of us ever thought to separate it from the genus Psalidognathus. I believe that Prionocalus [sic] Cacicus White belongs to the same genus, and that the remarkable differences pointed out by the author are only specific. In 1850, White described two more species in the genus: Prionacalus atys from Peru and Prionacalus iphis from Mexico. White (1850) recognized his mistake on the sex of the syntypes of P. cacicus:

4 THE COLEOPTERISTS BULLETIN 67(3), In the Annals and Magazine of Natural History, vol. XV. p. 108, I have described under the name of Prionacalus Cacicus, a curious genus from Mexico, allied to Psalidognathus, G. R. Gray. I regarded the two specimens as male and female of the same species, but it would seem that they are both males, and as they are considerably different, must be different species; what was deemed the male may retain the name Prionacalus Cacicus; it is figured on plate 8. fig. 1. of the above volume. TheotherspecimenmaybenamedPrionacalus Iphis; it is figured on plate 8.f.2. Thomson (1861) separated Psalidognathus from Prionocalus [sic], (translation): Mandibles less elongate; antennae spinose; prothorax subtriangular, anteriorly very wide and posteriorly narrow, lateral with 4 spines [two on each side]; humerus spinose; prosternum protruding, with apex elongate [prosternal process] and curved; mesosternum unarmed; hind legs very elongate; anterior tibiae of male dilated, inner margin longitudinally canaliculate, hairy; hind tarsi elongate Prionacalus; and Mandibles more elongate, curved, subvertical; antennae not spinose; eyes less separated; palpi very elongate; prothorax laterally with 4 spines, transverse, very short; humerus spinose; prosternal apex [prosternal process] very prolonged, curved; mesosternum protruding; anterior tibiae of male dilated, inner margin longitudinally canaliculated; hind tarsi elongate; female apterous Psalidognathus. Some of the characters described by Thomson (1861) are not useful for separating the two genera since they are either shared by both or highly variable (for example mandibular length). Thomson (1861) was the first to allocate Prionacalus and Psalidognathus in a single group (translation): Head broader; posterior area of head with a spine on each side; eyes dorsally apart; antennae elongate; mandibles inferiorly excavate, large, curved and subvertical; palpi very elongate; prothorax transverse, very short, laterally pluridentate; elytra in male a little wider and shortened; prosternum protruding; mesosternum sometimes unarmed; anterior tibiae dentate, in male, inwardly, distinctly longitudinally canaliculated; hind tarsi elongate; female apterous Psalidognathitae. This description is inaccurate with regard to Prionacalus, mainly because the prothorax is not strongly transverse and females of Psalidognathus are not apterous. Thomson (1864) redescribed his Psalidognathitae, retaining only Psalidognathus and Prionacalus (misspelled as Prionocalus) as members (translation): Head large, extended in a neck [elongate behind eyes]; head laterally [behind eyes] with a spine on each side; eye subfinely granulate; antennae in female shorter than body; mandibles large, curved, inferiorly excavate; palpi very elongate; prothorax transverse, very short, laterally pluridentate, but not crenulate; female with wide elytra sub-shortened; abdomen always manifestly shorter; anterior tibiae dentate, in male, inwardly, distinctly longitudinally canaliculated and dilated; hind tarsi very elongate; female wide, apterous, elytra shorter than abdomen. Actually, although the membranous wings are very reduced and small, they are present in both sexes. Although Lacordaire (1868) maintained Prionacalus (misspelled as Prionocalus) separate from Psalidognathus, he questioned its validity as a genus (translation, p. 41): Genus very near to Psalidognathus and contested [Reiche, 1850], for failing to recognize the essential character which distinguishes it, which is that in both sexes the metasternum is as in Psalidocoptus, that is to say, extremely short [note by Lacordaire However, I consider this character as provisional, because it is possible that this character existed in some Psalidognathus that I have not seen]. Taschenberg (1870) described Psalidognathus wallisi, based on two specimens from Ecuador (Loja). According to Lameere (1910), the two specimens comprised two distinct species (translation): I could see in the Museum of Halle [Martin- Luther-Universität, Wissenschaftsbereich Zoologie, Halle - Germany] that Taschenberg had joined a male of P. modestus Fries with a female of Prionocalus [sic] cacicus White to describe his species Psalidognathus Wallisi Waterhouse (1872) described another species from Ecuador, Prionacalus buckleyi, basedona male and female. The author also added a key to recognize the four species known at that time, and described for the first time the female of P. cacicus collected in Peru. Gemminger and Harold (1872) considered Prionacalus as valid, and correctly summarized the taxonomic history of the name of the genus. Lacordaire (1876) figured amaleofp. cacicus, but according to Lameere (1910), the specimen is a male of P. atys. Thomson (1877) published Waterhouse s (1872) key to Prionacalus in Latin, and affirmed that the genus (misspelled as Prionocalus) does not occur in Mexico (translation): It is incontestably by mistake that Mr. White and other authors had recorded Mexico as homeland of P. cacicus. The specimen of this species known by Mr. Reiche (Ann. Soc. Ent. France, 1850, p. 265) was taken in Ecuador by a traveler who gave it to Mr. Lemoinne, former Consul General of France in Lima. According to Mr. A. Lasèque (Musée botanique de Benjamin Delessert, Paris, 1845, p. 209), Mr. Hartweg, who also took the Prionocalus [sic] Cacicus, sent his latest collections to Mr. Delessert late 1843, from the area of Guayaquil, mountains around Loja, from the Andes of Quito, Popayau, and Bogota,

5 204 THE COLEOPTERISTS BULLETIN 67(3), 2013 from the edges of the Magdalena, and especially from Chimborazo, Artisana and Pichincha. Finally, Mr. Salle told us he had never found Prionocalus [sic] in Mexico. Lucas (1886) considered Prionocalus [sic] a valid genus and commented: I do not know if this observation has so far been reported by authors because White, to whom we owe the creation of this genus,ann.amag.ofnat.hist.,t.xv,p.109 (1845), does not mention these differences among the specific characters of the species included in this genus. I would add that the femurs of third pair of legs surpass the elytra and abdomen in individuals of various sizes that I observed, and the femurs of other pairs of legs are more or less swollen. Bates (1892) described Prionocalus [sic] whymperi from Ecuador, based on two males and one female, and Prionocalus [sic] trigonodes, also from Ecuador, based on a single male. Bates also commented on a single male specimen of P. buckleyi from Ecuador, examined by him, that differed from the typical form by having the elytra slightly less coarsely sculptured. Lameere (1910) considered P. whymperi as a synonym of P. buckleyi. Quentin and Villiers (1983) revalidated P. w h y m p e r i and placed P. trigonodes under synonymy of P. b u c k l e y i. Hüdepohl (1985) revalidated P. trigonodes (translation, p. 120): Since the revision of LAMEERE (1911), two species had been described as new: Psalidognathus (Prionocalus [sic]) woytkowskii HEYROVSKY, 1960, from Peru and Prionocalus [sic] demelti QUENTIN AND VILLIERS, 1982 from Colombia. According to the description and figures, woytkowskii HEYROVSKY, 1960 agrees completely with trigonodes BATES, 1891, the latter occurs not only in Ecuador, but also in Peru (Peru, Rio Huallagas, E. G. PEÑA leg., IV-1970, Serie in coll. m). Woytkowskii likely is a synonym of trigonodes BATES, Unfortunately, Hüdepohl (1985) did not formalize the synonymy, and makes his doubts clear in the key where he listed P. woytkowskii and P. trigonodes together in couplet 6. Gahan (1894) described Prionocalus [sic] gunteri from Ecuador, based on a single male specimen. Lameere (1910) synonymized that species with P. buckleyi, stating (translation): P. Gunteri Gahan was described on one male with abdomen finely punctate, elytra separately rounded apically, appendices reddish, with very prominent shoulder of elytra, and elytra less rough towards apex. Waterhouse (1900) described three new species from Ecuador (all misspelled as Prionocalus): P. simonsi, based on at least two males; P. whitei, based on a single male; and P. uniformis, based on two males. Lameere (1910) later synonymized P. whiteiwith P. buckleyi. Lameere (1901) considered Prionacalus (misspelled as Prionocalus) as part of Psalidognathus stating (translation): The Anacolines include: Cacosceles, Psalidocoptus, Micropsalis (and probably including Acalodegma, unknown to me), Psalidognathus (including Prionocalus) Kolbe (1902) described P. emmae, based on one male from Ecuador. Lameere (1910) placed this species under synonymy with P. buckleyi. Lameere (1910) considered Prionacalus (misspelled as Prionocalus) a subgenus of Psalidognathus (translation): These insects, always without metallic color, have all characters of the more inferior of the Psalidognathus, P. erythrocerus, and do not differ only by: 1, absence of membranous wings in both sexes, this has as consequence a pronounced shortening of metasternum in male, more than in female, and also a shortening of abdomen and elytra in male; 2, an elongation and strengthening of legs in male, character compensating for the loss of aerial locomotion; this particularity, combined with the shortening of thorax, results from the fact that the hind femora in male greatly exceed apex of body; 3, an enlargement of last segment of palp, more pronounced in male than in female. The sides of metasternum are glabrous in both sexes; the anterior edge of prothorax does not advance on sides of head; and the antennal segments are angular after antennomere III at outer and inner angles. Lameere (1910) also divided Psalidognathus (Prionocalus [sic]) into two groups. The first one was defined as follows (translation): The pronotum and elytra are covered with strong roughness. He included in this group: Psalidognathus iphis; Psalidognathus uniformis; Psalidognathus buckleyi; and Psalidognathus trigonodes. The second group was defined as follows (translation): The pronotum is finely rugose; the roughness of elytra is slightly marked, mainly apically. In this latter group, he included: Psalidognathus atys; Psalidognathus simonsi; andpsalidognathus cacicus. Lameere (1913, 1919) maintained Prionacalus (misspelled as Prionocalus) as a subgenus of Psalidognathus, and considered as valid the same species as in Lameere (1910). Gilmour (1954) followed Lameere (1910) in considering Prionocalus (misspelled as Prionocalus) as a subgenus of Psalidognathus. Heyrovsky (1960) described Psalidognathus (Prionocalus [sic]) woytkowskii based on three males and one female from Peru. Hüdepohl (1985) later considered P. woytkowskii as a synonym of P. trigonodes. Duffy (1960) described the larva of Prionacalus atys. Prionacalus was considered a subgenus of Psalidognathus and wrongly spelled as Prionocalus. Quentin and Villiers (1983) considered Prionacalus as distinct from Psalidognathus, and described Prionocalus [sic] demelti from Colombia, based on four males and two females. The authors gave

6 THE COLEOPTERISTS BULLETIN 67(3), a key to the species and considered as valid the following nine species: P. iphis; P. uniformis; P. demelti; P. woytkowskii; P. buckleyi; P. whymperi; P. simonsi; P. atys; andp. cacicus. Hüdepohl (1985) described the latest species in Prionacalus (misspelled as Prionocalus): Prionacalus giovannii. The species was described from Ecuador and based on two males. The author gave another key, considering the following nine species in Prionacalus: P. i p h i s ; P. uniformis; P. giovannii; P. trigonodes; P. buckleyi; P.simonsi; P. atys; P.cacicus; P. demelti. Prionacalus whymperi was omitted without any explanation. Komiya (2001) gave another key to Prionacalus (correctly spelled), and figured specimens of P. casicus [sic], P. w h y m p e r i, P. woytkowskii, P. iphis, and P. demelti. Monné (2006) listed the same species as in Hüdepohl (1985), but included Prionacalus whymperi. The generic name was correctly spelled. Jeniš (2010) considered the genera Prionacalus, Psalidognathus, and Apterocaulus Fairmaire, 1864 in the tribe Psalidognathini. However, this name has not been accepted by researchers working in Cerambycidae. For example, the name is not included in the checklist of Neotropical cerambycoid beetles (Bezark and Monné 2013) or the world catalogue of Cerambycidae (Tavakilian and Chevillotte 2013). Jeniš (2010) also published photographs of Prionacalus, some of which he labeled as type specimens. Unfortunately, these photographs were digitally modified, making it difficult or impossible to recognize them when compared to the actual type specimens. The following is a list of errors related to the photographs of Prionacalus in Jeniš (2010): 1) in at least one case, it is clear that the photograph does not match the actual type of the species (p. 19, syntype male of P. atys); 2) a female specimen was incorrectly figured as the type specimen of P. cacicus; 3) the terms holotype, syntype, and type were used inappropriately. In addition to these errors, P. gunteriand P. iphisare listed under the single name: P. iphis. However, Jeniš (2010) apparently considered these as distinct species. For example, the specimen figured on page 16 and labeled as a syntype of P. gunteri is the same specimen figured on page 122. The specimen figured does not match the holotype of P. gunteri (i.e., slender antennae, elytral apex laterally with a short tooth). We conclude that if the specimen figured is the actual type specimen deposited at the BMNH, then it was strongly modified digitally. Prionacalus White, 1845 Prionus (Prionacalus) White 1845: 109. Prionacalus; White 1853: 7; Gemminger and Harold 1872: 2754 (catalogue); Monné and Giesbert 1994: 14 (checklist); Monné 1995: 60 (catalogue), 2006: 82 (catalogue); Martínez 2000: 80; Di Iorio 2003: 2; Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Monné et al. 2012: 7 (checklist). Prionocalus; Chevrolat 1847: 469; Thomson 1861: 297, 331, 1864: 280, 468, 1877: 252, 260; Lacordaire 1868: 41; Waterhouse 1872: 261 (key to species); Quentin and Villiers 1983: 223, 225 (key to species); Hüdepohl 1985: 120 (key to species). Psalidognathus (Prionocalus); Lameere 1910: 374, 1913: 65 (catalogue), 1919: 121; Blackwelder 1946: 555 (checklist). Type Species. Prionus (Prionacalus) cacicus White, 1845 (monotypy). Redescription. Medium (ca. 24 mm) to large (ca. 74 mm) size, variable intraspecifically. Integument from slightly opaque to shining, brown to blackish brown, frequently with reddish parts. Male. Head: Prognathous, large (Figs. 16, 27, 40), excluding mandibles times longer than prothorax; very elongate behind eyes (distance from posterior ocular edge to prothorax times width of upper ocular lobe). Longitudinal dorsal exuvial cleavage line (coronal suture) usually not well-marked, often visible only behind eyes, occasionally absent. Vertex almost flat at base, gradually oblique towards middle of antennal tubercle, then abruptly becoming steeper (frequently distinctly vertical or bent inwards) towards clypeus (centrally usually less elevated when longitudinal furrow is deep) (this latter area would correspond to region of frons); coarsely, abundantly punctate, frequently also partially granulate, mainly laterally; usually with 2 cephalic carinae between upper ocular lobes, from antennal tubercle to about middle of area behind eyes (sometimes shorter), occasionally absent; with short, sparse setae. Area between cephalic carinae flat to distinctly depressed; frequently with shallow to deep, short (not reaching middle of eyes) to long (slightly surpassing posterior ocular edge) longitudinal furrow, occasionally almost absent. Antennal tubercle very distinct; apex rounded, turned sideward; bases of tubercles ranging from subcontiguous (separated only by the longitudinal furrow) to widely separated (distance between them about twice its width). Frontal sutures and frons usually indistinct. Epistomal suture usually laterally well-marked, often less so centrally (occasionally absent centrally). Clypeus with 2 areas: basal area subhorizontal, distal area not coplanar with basal area, usually from strongly oblique to vertical, in some species distinctly concave. Basal portion of clypeus narrow (centrally times width between tentorial pits), flat, centrally on basal 2/3 usually slightly

7 206 THE COLEOPTERISTS BULLETIN 67(3), 2013 obliquely elevated (occasionally not elevated, but with 2 gibbosities); glabrous or with sparse, very short setae; sculpture variable: coarsely, shallowly, abundantly punctate, or moderately finely, sparsely punctate, or almost smooth; anterior edge widely emarginate. Distal portion of clypeus smooth (sometimes with very sparse fine punctures), shining, glabrous. Labrum from narrow (length about 0.25 times width) to wide (length about 0.40 times width); at least basal area oblique in relation to distal region of clypeus; frequently strongly depressed centrally on distal area; distal area coplanar or not with basal area; distal edge straight or slightly convex or, more rarely, slightly concave (sometimes centrally feebly projected); surface shining, smooth, glabrous (sometimes with long, sparse setae); distal edge centrally with long setae (occasionally along nearly entire edge). Anterior tentorial pits well-marked. Eyes narrow (width equal to about 0.3 times length), finely faceted, distinctly emarginate, moderately small; ocular lobes with same width; upper ocular lobe a little shorter than lower ocular lobe; distance between upper ocular lobes times longest width of 1 lobe; apex of lower ocular lobes reaching level of middle of mandible at base. Lateral area of head, behind eyes, with 1 tubercle, intraspecifically variable in size (from moderately small to large) and shape (from distinctly acute at apex to rounded); surface usually roughened, with short, sparse setae between level of tubercle and prothorax (occasionally, with some sparse setae between level of tubercle and eyes). Genal apex usually with downward projection (small or absent in some species; in others always very distinct). Gula trapezoidal (occasionally, length equal to about 0.4 of total length of ventral surface); with short, abundant setae on base, sparser towards apex. Posterior tentorial pits well-marked. Gular sutures well-marked, deep. Submentum moderately well-delimited (sometimes distinctly delimited), usually depressed from gula to near anterior 1/3 where distinctly elevated; sculpture variable; with short, sparse setae (often almost glabrous). Mandible intraspecifically variable in size and shape (variation usually linked to size of specimen), but always long and apically curved downward and inward; left mandible different in form from right one; coarsely, abundantly punctate, with short sparse setae (except on apical portion). Mentum transverse (longest length about 0.35 times width); anterior margin trilobed (central lobe occasionally very short or absent); about as wide as 1/2 width of submentum at anterior margin; coarsely, moderately abundantly punctate (sometimes only laterally); with long, moderately sparse setae. Galea very short, reaching from about middle to distal 1/3 of palpifer, with long, abundant setae turned upwards. Palpifer, with distal membranous area not extended, about as long as maxillary palpomere I. Maxillary palp very long ( times anterior width of head); palpomere I shorter than remaining segments, II longer than III, and IV longer than others; palpomere IV strongly enlarged towards apex. Ligula bilobed, each lobe obliquely projected forward. Labial palp very long ( times anterior width of head); palpomere I times length of II, and palpomere III slightly longer than II; palpomere III strongly enlarged towards apex. Antennae with 11 segments; almost reaching to distinctly surpassing elytral apex; scape conical or slightly enlarged towards apex, distinctly surpassing posterior ocular edge, shorter than antennomere III, at least dorsally coarsely punctate; pedicel very small (smaller than 0.2 times length of antennomere III) to small (slightly longer than 0.2 times length of antennomere III); antennomere III longest, enlarged towards apex, ventral surface with or without sensorial area near apex (if present, not divided by carina); ventral surface of antennomere IV with sensorial area on distal 1/2, divided or not by carina, and sometimes with small dorsal sensorial area; ventral surface of antennomere V almost entirely occupied by sensorial area, always divided by at least 1 carina (frequently more), dividing sensorial area into irregular areas, and sometimes with small dorsal sensorial area; ventral surface of antennomeres VI-XI entirely occupied by sensorial area, always divided by longitudinal carina, and dorsally with sensorial area (larger and divided by carinae from VII or VIII); basal antennomeres with short, sparse setae (usually only at apex), and distal antennomeres glabrous (occasionally, antennomere XI with very short, sparse setae on apex). Thorax: Prothorax transverse (longest width times central length); latero-anterior angle slightly projected forwards, latero-posterior angle from projected sideward (frequently almost as a spine) to rounded; lateral edges with spines (sometimes almost coarsely crenulate); anterior edge from straight to somewhat rounded (not rarely centrally emarginate); posterior edge from slightly sinuate to rounded. Pronotum convex, centrally depressed, and laterally expanded; surface coarsely rugopunctate; centrally glabrous (occasionally with some setae), and laterally with short, sparse setae. Prosternum convex, strongly elevated centrally; surface usually striate (sometimes coarsely) and finely, sparsely punctate; with short, sparse setae (sometimes almost glabrous). Prosternal suture distinct, straight from procoxal cavity towards apex. Procoxal cavities widely opened posteriorly. Prosternal process long, reaching basal 1/3 to middle of mesocoxae. Mesosternum narrow, separated by complete sutures from mesepisterna. Mesocoxal cavities laterally open.

8 THE COLEOPTERISTS BULLETIN 67(3), Mesepisternum with or without short setae. Mesosternal process deeply sulcate (sometimes less so), narrowed towards apex. Metasternum reduced; surface moderately finely punctate to coarsely punctate; with short, sparse setae. Metepisternuma with margins parallel at basal 3/4 and convergent at distal 1/4 (inner margin more distinctly), or slightly curved at inner margin, mainly at distal 1/2; surface abundantly or moderately coarsely punctate, with short, sparse setae. Scutellum short, transverse, rounded or moderately straight distally, and emarginate or not at center of this edge (variable intraspecifically); surface from finely to coarsely punctate (variable intraspecifically), and short and moderately abundantly setose or almost glabrous (intraspecifically variable). Elytra fused at suture; outer margin parallel at basal 1/3 or 1/2, distinctly convergent in remainder; humeral angle situated below level of scutellar apex, usually strongly spiniform; elytral surface, behind humeral spine, usually abruptly sloping downward; apices usually individually rounded (occasionally almost truncate, and sutural angle sometimes projected); glabrous or with very short and sparse setae at base; surface entirely coarsely rugose (somewhat vermiculate) or finer rugose on distal 2/3; membranous wings (Fig. 36) very small (about 0.2 times length of elytron). Abdomen: Occasionally surpassing elytral apex, narrowed from base to apex, strongly convex at ventrite I, gradually becoming flatter towards apex; ventrite I longest, II slightly longer than III, III and IV with subequal length, V distinctly emarginate at apex. Median lobe of genitalia times length of tegmen; ventral lobe of distal portion of median lobe distinctly narrowed towards apex, not surpassing apex of dorsal lobe; dorsal lobe of distal portion of median lobe distinctly emarginate at apex; basal apophysis of median lobe enlarged towards apex, times total length of median lobe; lateral lobes of tegmen times total length of tegmen, with long, dense setae on dorsal distal 1/2 (occasionally only at distal 1/3), distinctly sparser on basal 1/2 (occasionally only at basal 1/3), ventral surface and lateral margins with long, dense setae. Legs: Trochanter with elliptical depression ventrally, coarsely, sparsely punctate (sometimes abundantly punctate), with long, sparse setae (sometimes shorter or more abundant). Femora narrower dorsally than ventrally, dorsally convex, laterally oblique, and ventrally sulcate; pro- and mesofemora subequal in length, metafemora distinctly longer; lateral surface transversely rugose and punctate (variable in shape and concentration), usually coarsely on profemora; apex of metafemora usually surpassing elytral apex (sometimes only reaching elytral apex). Protibiae distinctly swollen centrally, narrow at base, apex somewhat narrower than middle area, distinctly compressed dorso-ventrally on distal 1/5 or 1/6; latero-distal surface with moderately abundant denticles (sometimes with distinct spines on margin), usually with distinct depression near apex, ventral surface longitudinally deeply sulcate (length of depression variable), with dense setae in depression; apex with 2 spurs (the lower is longest). Mesoand metatibia enlarged from base to apex, usually with small spines and denticles on proximal surface (sometimes absent on metatibiae or both), depressed on distal 1/3 or 1/4 of proximal surface (depression occasionally distinct from basal 1/3, but always more conspicuous distally; sometimes almost inconspicuous); apex with 2 spurs, inner most larger. Tarsi long, mainly meso- and metatarsi; protarsi moderately enlarged (occasionally slightly more than mesotarsi); mesotarsi slender and longer than protarsi, and metatarsi slender and longer than mesotarsi; metatarsomeres I-III usually spined at apices (less distinctly on protarsomeres). Female. The main differences with males are as follows: mandibles shorter than head; last segment of palpi (maxillary and labial) less expanded towards apex (usually times longest width of male); antennae shorter, never reaching elytral apex; elytra usually more rounded from base to apex; elytra sometimes not fused at suture; protibiae ventrally not sulcate, not swollen centrally; apex of metafemora not or rarely reaching elytral apex; abdomen usually surpassing elytral apex; apex of ventrite V from truncate to slightly emarginate (sometimes rounded). Included Species. Prionacalus atys White, 1850; Prionacalus cacicus (White, 1845); Prionacalus demelti Quentin and Villiers, 1983; Prionacalus iphis White, 1850; Prionacalus uniformis Waterhouse, 1900; Prionacalus whymperi Bates, 1892; and Prionacalus woytkowskii (Heyrovsky, 1960). Geographical Distribution. Ecuador, Peru, Colombia, Bolivia, and Argentina. Diagnosis. Prionacalus is similar to Psalidognathus. Males differ as follows: length of head (excluding mandible) + prothorax times elytral length ( times in Psalidognathus); brachypterous (membranous wings well-developed in Psalidognathus); metasternum distinctly shortened (not shortened in Psalidognathus). Females differ as follows: body, in lateral view, wider (body narrower in lateral view in Psalidognathus); two anterior spiniform projections on lateral margins of prothorax usually broadly joined and occasionally strongly protracted (two anterior spiniform projections on lateral margins of prothorax distinctly separated and more strongly protracted in Psalidognathus). Prionacalus differs from Apterocaulus by the body less flattened dorsoventrally, the dorsal surface of the head usually with two carinae, the mid-lateral spiniform projection of

9 208 THE COLEOPTERISTS BULLETIN 67(3), 2013 the prothorax shorter and not strongly curved backwards, and the distinctly narrower tarsi. In Apterocaulus, the body is flatter dorso-ventrally, the dorsal surface of is head lacks carinae, the mid-lateral spiniform projection of the prothorax is distinctly longer and strongly curved backwards, and the tarsi are distinctly shorter and wider. Remarks. All species of Prionacalus exhibit notable allometry in the proportions of the head, size and shape of the mandible, palpi, antennae, and elytra. The considerable number of specimens with features intermediate among the species suggests that hybridism may occur. A phylogenetic analysis including both morphological and molecular data is needed to test this hypothesis. KEY TO THE SPECIES OF PRIONACALUS 1. Elytral sculpture different on basal and distal halves Elytral sculpture uniform (or similar) throughout Males: Last maxillary and labial palpomere notably enlarged at apex (longest width equal to about 0.65 times its length); cephalic carinae (Fig. 18) not notably elevated at apex. Females: Cephalic carinae slightly elevated (Fig. 23); antennae reaching about apex of basal 1/3 of elytra (sometimes almost elytral middle) (Fig. 23). Ecuador, Peru......Prionacalus atys 2. Males: Last maxillary and labial palpomere slender towards apex (longest width times its length); cephalic carinae (Fig. 29) distinctly elevated at apex. Females: Cephalic carinae distinctly elevated (Fig. 37); antennae reaching base of distal 1/3 1/4 of elytra (Fig. 37). Ecuador, Colombia, Peru Prionacalus cacicus 3. Genal apex with long, acute projection (Fig. 42). Colombia...Prionacalus demelti 3. Genal apex with short, acute projection or rounded and not projected (Figs. 53, 66, 78) Prosternal process with spiniform projection on underside of apex (Fig. 14); protibia of male enlarged near middle. Ecuador......Prionacalus whymperi 4. Prosternal process without spiniform projection on underside of apex or with rounded projection (Fig. 15); protibiae of male gradually enlarged from base Cephalic carinae almost indistinct in both sexes (Figs. 66, 72); sulcus between cephalic carinae distinctly deep (Fig. 66). Argentina, Bolivia, Peru... Prionacalus woytkowskii 5. Cephalic carinae distinct, mainly in males (Figs. 78, 84, 124); sulcus between cephalic carinae slightly or moderately deep (Figs. 78, 84, 124) Antennae of male not reaching elytral apex (Fig. 124). Female unknown. Peru......Prionacalus uniformis 6. Antennae reaching or surpassing elytral apex in male (Fig. 76), and distal 1/3 in female (Fig. 84). Ecuador, Peru...Prionacalus iphis Prionacalus atys White, 1850 (Figs. 8, 16 26, , ) Prionacalus Atys White 1850: 11, pl. 13, fig. 4; 1851: 70, 71, 1853: 8; Gemminger and Harold 1872: 2754 (catalogue). Prionacalus atys; Waterhouse 1880: 485; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 82 (catalogue), 2012: 7 (checklist); Di Iorio 2003: 3 (distribution); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 19 (syntypes male and female), 118 (male), 119 (female). Prionocalus Atys; Lacordaire 1868: 42 (note); Waterhouse 1872: 261 (key); Thomson 1877: 260 (key). Psalidognathus (Prionocalus) Atys; Lameere 1910: 377, 1913: 66 (catalogue), 1919: 121. Psalidognathus (Prionocalus) atys; Blackwelder 1946: 555 (checklist); Duffy 1960: 15, 69 (larva). Prionocalus atys; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121; Komiya 2001: 32 (key). Prionocalus Simonsi Waterhouse 1900: 504. New synonymy. Psalidognathus (Prionocalus) Simonsi; Lameere 1910: 378, 383 (key), 1913: 66 (cat.), 1919: 121. Psalidognathus (Prionocalus) simonsi; Blackwelder 1946: 555 (checklist). Prionocalus simonsi; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121 (key). Prionacalus simonsi; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 61 (catalogue), 2006: 83 (catalogue); Di Iorio 2003: 3 (distribution); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2008: 95 (male), 2010: 18 (males and females), 120 (male), 121 (female), 145 (syntype male). Redescription. Male (Fig. 16). Head and mandibles blackish (occasionally, head dark brown); maxillary and labial palpi reddish; antennae gradually lighter towards apex, usually scape, pedicel, and basal 2/3 of antennomere III blackish, distal

10 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus species. 1) P. cacicus, male, maxillary palpus; 2) P. cacicus, male, maxillary palpus; 3) P. woytkowskii, male, maxillary palpus; 4 5) P. iphis, male, maxillary palpus; 6) P. whymperi, male, maxillary palpus; 7) P. demelti, male, maxillary palpus; 8) P. atys, male, maxillary palpus; 9 10) P. woytkowskii, male, antenna; 11 12) P. cacicus, male, antenna; 13) P. demelti, male, antenna; 14) P. whymperi, female, prosternal process, lateral view; 15) P. iphis, female, prosternal process, lateral view.

11 210 THE COLEOPTERISTS BULLETIN 67(3), /3 of antennomere III, and basal 2/3 of IV and V dark brown, distal 1/3 of antennomeres IV and V reddish, antennomeres VI-XI reddish; pronotum blackish (sometimes dark brown); ventral surface of thorax dark brown (usually, prosternum almost blackish); elytra dark brown on basal 1/3, gradually reddish towards apex; ventrites I-IV dark brown, usually distal margin blackish, ventrite V darkbrown on base, reddish towards apex (occasionally entirely reddish); profemora dark brown; meso- and metafemora reddish brown, with some parts dark brown; protibiae dark brown on basal 1/3, gradually reddish towards apex; meso- and metatibiae dark brown on base (sometimes on basal 1/3) reddish on remaining surface; tarsi reddish, except apex of tarsomere V and claws. Head: Coarsely punctate-vermiculate between cephalic carinae, coarser on 1/2 near clypeus; area between apex of cephalic carinae and pronotum coarsely, abundantly, confluently punctate; area on 1/2 near clypeus with longitudinal, well-marked furrow (occasionally present on area between antennal tubercles); distal margin of vertex (close to abrupt declivity) centrally distinctly emarginate. Antennal tubercle moderately coarsely, sparsely punctate on base (sometimes with confluent punctures), gradually becoming smoother towards apex; distinctly far from each other (distance between them equal to its width, or slightly larger). Cephalic carinae (Fig. 18) very distinct, elevated from base to apex; convergent from antennal tubercle to little after eyes, then divergent towards apex; apex not or slightly elevated, usually not forming triangular tubercle. Area behind eyes variable: not finely striate, with abundant small asperities, mainly between eyes and dorsal surface of lateral tubercle; finely striate, with abundant asperities only behind lower ocular lobes; finely striate, with sparse asperities only behind lower ocular lobes. Lateral tubercle of head moderately large, rounded (Fig. 16) to distinctly acute (Fig. 121) at apex. Subhorizontal area of clypeus finely, moderately sparsely punctate, more distinctly basally. Distal area of clypeus distinctly concave. Labrum distinctly concave. Distance between upper ocular lobes times longest width of each lobe. Gena rounded at apex (Fig. 18), without downward projection. Submentum laterally shallowly, coarsely, abundantly punctate, centrally finely, abundantly punctate or moderately finely, transversely striate (occasionally entire surface of submentum transversely striate, stria can be moderately coarse). Mandible (Fig. 19) times head length; inner margin laminar; surface coarsely, abundantly punctate, gradually finer towards apex (smooth near inner margin); left mandible with 2 distinct teeth near base (sometimes small and together protracted); right mandible with 2 small, protracted teeth near base and another large tooth close to base of distal plate; distal plate of left mandible wider than that of right mandible (in one specimen examined, right mandible identical to left). Longest width of last maxillary (Fig. 8) and labial palpomeres equal to about 0.65 times its length. Antennae not reaching distal 1/3 or 1/4 of elytra; scape distinctly enlarged from base to apex; antennomeres III-IV usually rounded at apex on both sides (sometimes slightly projected at outer distal angle); antennomeres V-X usually distinctly dentate at outer distal angle. Thorax: Lateroanterior angles of prothorax slightly projected forward, rounded (occasionally acute); lateroposterioranglesofprothoraxfromacuteanddistinctly projected sideward (tooth-like) to slightly or not projected (in both cases, rounded); anterior 2 teeth of lateral margin protracted, with anterior tooth smallest (sometimes absent or almost so); latero-basal tooth large, usually slightly curved upwards. Pronotum with distinct median depression and another smaller depression laterally on each side; coarsely, confluently punctate on disc, laterally with coarser and sparser punctures, but with smooth, shiny areas between punctate areas (general appearance of lateral area much more shiny than central area). Elytra moderately coarsely vermiculate on central subtriangular area (larger area on elytral base) from base to middle (sometimes up to base of distal 1/3); remaining surface distinctly finer sculptured; times longer than prothorax; apex individually rounded. Apical tooth of humeral projection absent or distinct. Abdomen: Not surpassing elytral apex. Ventrites I-III finely, sparsely punctate (Fig. 17); ventrite IV finely, abundantly punctate; ventrite V coarser, distinctly, more abundantly punctate. Legs: Apex of metafemora surpasses elytral apex by about basal 1/3 2/5. Ventral sulcus of protibiae distinct from near base to distal constriction. Protarsomeres I and II (Fig. 20) short, moderately wide; protarsomeres I-III with very small denticle at apex; mesotarsi shorter than metatarsi (Fig. 21); metatarsi (Fig. 22) not notably slender. Female (Fig. 23). Color as in males, with same variations, except abdomen entirely dark brown with apex blackish (in one female the body is almost entirely reddish probably a teneral specimen). Inner margin of mandibles (Fig. 25) laminar; inner margin as in males, but teeth smaller. Antennae reaching about apex of basal 1/3 of elytra (sometimes almost mid-elytra). Longest width of last maxillary and labial palpomere about 0.4 times length. Cephalic carinae as in males, but less elevated or almost absent, and often less divergent distally. Distance between upper ocular lobes times longest width of 1 lobe. Lateral tubercle of head small, acute at apex. Prothorax as in male. Elytra times longer than prothorax,

12 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus atys. 16) Male, dorsal view; 17) Male, ventral view; 18) Male, head, dorso-lateral view; 19) Male, mandibles; 20) Male, protarsus; 21) Male, mesotarsus; 22) Female, metatarsus; 23) Female, dorsal view; 24) Female, ventral view; 25) Female, mandibles, 26) Female, metatarsus.

13 212 THE COLEOPTERISTS BULLETIN 67(3), 2013 elytral sculpture as in male or, rarely, uniform throughout. Abdomen distinctly surpassing elytral apex. All ventrites (Fig. 24) finely, sparsely punctate. Meso- and metatarsi (Fig. 26) shorter than in male. Dimensions. Male/female. Total length (including mandibles) = / mm; prothoracic length = / mm; prothoracic width between apices of anterior angles = / mm; prothoracic width between apices of posterior angles = / mm; humeral width = / mm; elytral length = / mm. Geographical Distribution. Ecuador (Waterhouse 1900), Peru (White 1850). (Mexico is not considered a country where this species occurs). Material Examined. ECUADOR: (no other data), 1 male, 2 females (ZMHB); male (no other data) (MPCO). Loja: female (no other data) (IRSN). Manabí: Portoviejo, female, (ex Nonfried Collection) (IRSN). Azuay: Cuenca, female, I.1971 [no collector indicated] (MPCO). PERU: 1 male, 1 female, (ex Desbrochers Collection; no other data) (IRSN). Additionally, we examined specimens with the following label: MEXICO, male, (ex Caulle Collection; no other data) (IRSN); female, (ex E. A. Klages Collection; no other data) (EMEC); male (no other data) (EMEC). Types and Type Localities. Of Prionacalus atys (Figs ): Described from Andes of Peru, apparently, based on a single male, deposited in the BMNH. The descriptions in White (1850, 1851) do not suggest that he had a female of the species. The BMNH website (2013) records one syntype of Prinocalus [sic] atys. However, Bezark (2013) figured two syntypes in this collection: one male and one female (both photographed by Henry Hespenheide). Of Prionocalus [sic] simonsi (Figs ): Described based on males (unknown number) from Ecuador ( in wood west of Cuença [sic], 2600 m ). Cuenca is a city in Azuay Province. According to Chubb (1919): The following notes are based, chiefly, on a collection made by the late Perry O. Simons in the Andean regions of Ecuador, Peru, Bolivia, and Argentina from the latter part of 1898 to November 1901, at varying altitudes up to 5000 metres The expedition was a private undertaking initiated and financed by Mr. Oldfield Thomas, F.R.S., of the Department of Zoology, British Museum (Natural History), whose enthusiasm is so well known among mammalogists and who has done so much to advance that branch of Zoological Science. His object was to obtain a collection, as complete as possible, of the mammals of the northern portion of the South American Andes, but, with his usual generosity, he allowed Simons to collect birds also during his journey. The specimens thus collected by Simons were acquired by the British Museum and form a particularly welcome addition to the Bird-Room We do not know if all specimens of Coleoptera collected by Perry O. Simons are deposited in the BMNH. The website of the BMNH (2013) records only one syntype of Prinocalus [sic] simonsi. EHN recorded that there is only one male type deposited at BMNH. It is possible that the indication of more than one measure in the original description was a mistake: Long mm. Remarks. According to Waterhouse (1900), This species [P. simonsi] closely resembles P. atys, White, in form, colour, and sculpture, but is at once distinguished by the posterior angles of the thorax being slightly rounded instead of acute and slightly projecting, as they are in P. atys. The head is coarsely and closely rugose, with a short, not very acute, conical tubercle behind the eye. The mandibles are shorter than the head. The thorax is less rugose than the head, and the scape on each side of the disk, although rather closely punctured, is smoother. The elytra are one-third longer than broad, much narrowed towards the apex, vermiculate-rugose at the base, the sides and apex (which are impressed) with much finer rugose sculpture; the apex of each elytron is rounded. The femora are more or less pitchy, the tibiae almost entirely so, the tarsi rather paler. Lameere (1910) wrote of P. simonsi (translation): It does not differ from P. A t y s except by the posterior angles of prothorax completely rounded. And in his key to the species, he separated those species by (translation): Posterior angles of prothorax projecting. Peru P. Atys. / Posterior angles of prothorax rounded. Ecuador P. Simonsi. Quentin and Villiers (1983) followed the same difference in their key (translation): Posterior angles of pronotum projecting. 8 [following to P. a t y s and P. cacicus] / Posterior angles of pronotum rounded P. simonsi Waterh. Finally, according to the key in Hüdepohl (1985) (translation): Lateral margin of pronotum with a small tooth at base (Peru, Ecuador) atys WHITE, 1845 / Lateral margin of pronotum rounded at base (Ecuador) simonsi WATERHOUSE, In summary, the authors who treated P. simonsi after the original description only repeated what was said by Waterhouse (1900). The only difference pointed out by Waterhouse (1900), the presence or absence of a tooth at the latero-posterior angles of the prothorax, is a variable character. We have examined males with teeth from very distinct (corresponding to P. a t y s )toalmostabsent(corresponding to P. s i m o n s i ). As there are no other differences between both species, we conclude

14 THE COLEOPTERISTS BULLETIN 67(3), that P. s i m o n s i is a variation of P. a t y s, and thus propose its synonymy. This can be confirmed by comparing photographs of syntypes of those species (Figs. 103 (P. a t y s ), 12 (P. s i m o n s i )) that show very similar heads, antennae, pronota, elytra, and legs. Prionacalus cacicus (White, 1845) (Figs. 1, 2, 11, 12, 27 39, , ) Prionus (Prionacalus) Cacicus White 1845: 110, pl. 8, figs. 1, 2. Prionus (Prionacalus) cacicus; Monné 2012: 132. Prionacalus Cacicus; White 1850: 10, 1851: 70, 1853: 7. Prionocalus Cacicus; Thomson 1864: 281, 1877: 260 (type locality, key). Psalidognathus (Prionocalus) cacicus; Reiche 1850b: 265; Lameere 1910: 378, 1913: 66 (catalogue), 1919: 121; Blackwelder 1946: 555 (checklist); Gilmour 1954: 43, pl. 8, fig. 3, pl. 9. Prionocalus cacicus; Lacordaire 1868: 42 (note); Waterhouse 1872: 261 (key), 262, 1900: 503; Lucas 1886: clxxvii; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121 (key); Pardo- Locarno 2006: 241 (fig. 3). Prionacalus cacicus; Gemminger and Harold 1872: 2755 (catalogue); Waterhouse 1880: 485; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 83 (catalogue); Verdcourt 2000: 18; Di Iorio 2003: 3 (distribution); Arai 2005: 34, fig. 1; Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 18 (8 figures, males and females), 19 (5 figures, males and females), 35, 112 (male), 113 (female); Monné et al. 2012: 7 (checklist). Prionacalus casicus [sic]; Komiya 2001: 29, 30 (fig. 1), 32 (key). Psalidognathus cacicus; Reiche 1850a: 249. Psalidognathus Wallisi Taschenberg 1870: 191 (female); Lameere 1910: 378 (synonymy). Prionacalus atys (not White, 1850); Lacordaire 1876: pl. 81, figs. 1, 1a; Lameere 1910: 378 (synonymy). Prionacalus buckleyi; Lameere 1919: pl. 6, fig. 2. Prionocalus Gunteri Gahan 1894: 221; Lameere 1910: 376 (synonymy). New synonymy. Prionacalus gunteri; Jeniš 2010: 16 (syntype male). Psalidognathus (Prionocalus) Buckleyi var. Gunteri; Lameere 1913: 65 (revalidation). Redescription. Male (Fig. 27). Head and mandibles blackish (occasionally with anterior margin of submentum reddish); maxillary and labial palpi reddish; antennae gradually lighter towards apex, usually scape and pedicel dark brown (sometimes slightly reddish); antennomere III brown (from dark to lighter); apically reddish from antennomere V or VI; pronotum blackish; ventral surface of thorax dark brown; elytra dark brown (usually blackish at base and, commonly, reddish brown towards apex); ventrite I dark brown (sometimes lighter), ventrite V reddish (frequently almost orange; occasionally dark brown), and intermediate ventrites gradually lighter (often distal margin of ventrites II-IV darker; occasionally entirely dark brown); legs reddish, partly dark brown (sometimes femora entirely blackish), except claws which are reddish on base and blackish on the remainder (occasionally, all tibiae bicolored: dark brown, at least on basal 2/3, and reddish on remainder). Head: Coarsely punctatevermiculate between cephalic carinae, especially apically; area between apex of cephalic carinae and pronotum coarsely, abundantly, confluently punctate (occasionally somewhat vermiculate); area on 1/2 near clypeus with longitudinal, well-marked furrow; distal margin of vertex (close to abrupt declivity) centrally, distinctly emarginate (sometimes emargination only is indicated). Antennal tubercles coarsely punctate near base, gradually smoother towards apex; distinctly separated (distance between them about twice their width). Cephalic carinae (Fig. 29) very distinct, elevated from base to apex but rarely less prominent; slightly convergent from antennal tubercle to a little after eyes, then divergent towards apex; apex distinctly elevated, forming a triangular tubercle; usually dorsally impunctate (sometimes, distinctly punctate). Area behind eyes with abundant, small asperities, usually closer towards dorsal surface of lateral tubercle, and less distinct towards prothorax. Lateral tubercle of head small and distinctly acute at apex (usually in small specimens) to large (Fig. 27) and rounded at apex (usually in large specimens). Subhorizontal area of clypeus usually moderately coarsely punctate, mainly basally (occasionally punctures fine and sparse, or coarse and shallow). Distal area of clypeus distinctly concave. Labrum not strongly concave. Distance between upper ocular lobes times longest width of 1 lobe. Gena rounded at apex (Fig. 29), rarely with weak downward projection. Submentum laterally, shallowly, abundantly punctate, centrally transversely sulcate (sometimes also sparsely punctate centrally; in large specimens, grooves are deep). Mandible in small specimens (Fig. 31) usually, at most, as long as head; inner margin laminar; surface coarsely, abundantly punctate, gradually finer and sparser towards apex (smooth near inner margin); left mandible with distinct, subtriangluar, large tooth between base and distal plate; right mandible with subtriangular tooth between base and distal plate (usually less distinct), and another close to base of distal plate; distal plate of left mandible wider than that of right mandible. Mandibles in medium sized specimens

15 214 THE COLEOPTERISTS BULLETIN 67(3), 2013 (Fig. 30) very variable: as long as head to longer, distinctly laminar to distinctly subcylindrical at basal 1/3; area between base and base of distal plate with 1 2 teeth (on one or both mandibles), and the tooth/teeth placed at middle or closer to base of distal plate. Mandibles in large specimens longer than head; basal 1/2 subcylindrical, gradually more laminar towards base of distal plate; left mandible with one tooth close to the base of distal plate; right mandible with distinct, large, triangular tooth close to base of distal plate. Longest width of last maxillary and labial palpomeres 0.35 (Fig. 2) to 0.45 (Fig. 1) times length. Antennae always surpassing elytral apex, usually by last 2 antennomeres, but occasionally only by 1 antennomere (depending, mainly, on elytral length); scape distinctly enlarged from base to apex; antennomeres III-X usually distinctly dentate at apex on both sides (sometimes only at outer distal angle on basal antennomeres), with variable width: narrow (Fig. 12) or thick (Fig. 11). Thorax: Latero-anterior angles of prothorax usually acute (sometimes rounded), projected forwards; latero-posterior angles of prothorax distinctly projected sideward, tooth-like (rarely slightly projected and rounded at apex); anterior 2 teeth of lateral margin strongly protracted, anterior tooth smallest; latero-basal tooth large, usually slightly curved upwards. Pronotum longitudinally depressed centrally; coarsely, confluently punctate (sometimes somewhat vermiculate), sometimes almost smooth on depressed area and basal parts; laterally coarsely, confluently punctate (occasionally sparsely punctate), but smooth and shiny between punctate areas. Elytra moderately coarsely vermiculate on basal 1/3 (sometimes only on basal 1/4), gradually distinctly finer towards apex; times longer than prothorax; apices usually individually rounded (occasionally rounded together or with sutural angle distinctly toothlike). Apical tooth humeral projection absent to very distinct. Abdomen: Usually not surpassing elytra. Ventrites I-III (Fig. 28) finely, sparsely punctate (in very large specimens, punctures more abundant, and somewhat coarser); ventrites IV-V slightly coarser, distinctly more abundantly punctate. Legs: Apex of metafemora usually surpassing elytral apex by 1/2, sometimes by about 1/3 (depending on elytral length). Ventral sulcus of protibiae (Fig. 32) distinct from near base to distal constriction. Protarsomeres I and II (Fig. 33) short, moderately wide; protarsomeres I-III with very small denticle at apex; mesotarsi shorter than metatarsi (Fig. 35); metatarsi normal or slender andlong(fig.34). Female (Fig. 37). Color as in males, with the same variations, except the abdomen which is entirely dark brown with the apex blackish. Inner margin of mandibles laminar; inner margin as in small males. Antennae reaching distal 1/3 or 1/4 of elytra. Longest width of last maxillary and labial palpomeres times length. Cephalic carinae as in males, usually with apex less elevated. Distance between upper ocular lobes longest width of 1 lobe. Lateral tubercle of head as in male and indicating similar variation. Prothorax as in males. Elytra times longer than prothorax. Abdomen usually distinctly surpassing elytral apex (occasionally not surpassing). All ventrites (Fig. 38) finely, sparsely punctate. Meso- and metatarsi (Fig. 39) shorter than in male. Dimensions. Male/female. Total length (including mandibles) = / mm; prothoracic length = / mm; prothoracic width between apices of lateral spines = / mm; humeral width (at humeral projection) = / mm; elytral length = / mm. Geographical Distribution. Ecuador (Reiche 1850b), Peru (Waterhouse 1872), Colombia (Jeniš 2010). (Mexico is not considered a country where this species occurs). Material Examined. ECUADOR: 1 male, 1 female (ex Moffarts Collection; no other data) (IRSN); 2 males (no other data) (ZMHB); male, Deyrolle col. (no other data) (ZMHB); male, 1907, C. Felsche col. (SNSD). Azuay: Cuenca, male, I.1971, [no collector indicated] (ZKCO). Loja: Utuana, male, 30.III.2000, F. Hovore col. (EMEC); Loja (no other data), female (IRSN); 3 males, 2 females, Abbé Gaujon col. (no other data) (IRSN); 2 males, de Mathan col. (ex Moffarts Collection; no other data) (IRSN); male, de Mathan col. (no other data) (IRSN); male, Abbé Gaujon col. (ex Moffarts Collection; no other data) (IRSN); male, Abbé Gaujon col., (no other data) (SNSD); male, , F. Ohaus col. (ZMHB); (2200 m), male, 2.VIII.1905, F. Ohaus col. (ZMHB); male (no other data), E. Witts col. (ZMHB); near Loja, female (ex Moffarts Collection; no other data) (IRSN); Cerro Villanaco (ca. 7 km W of the city of Loja; 3000 m), 2 males, 11.X.1905, Ohaus col. (ZMHB). Sucumbíos: La Bonita, male, X.1997, local collector (MPCO). Azuay: Cuenca, male, II.1975, local collector (MPCO). Tarqui (probably in Azuay province), male, 07.III.1965, Peña col. (MZSP). COLOMBIA: 1 male, 1 female (ex Caulle Collection; no other data) (IRSN). Huila: male, 1975 (no other data) (DHPC). Pichincha: Santa Inés, male, R. Haensch col. (SNSD). PERU: 3 males (ex Desbrochers Collection; no other data) (IRSN); mountain (no other data), Murray col. (ZMHB). Piura: Ayabaca, 2 males, 1 female, 20.III.2008, local collector (MPCO); 1 male, 1 female, II.2008, (no collector indicated) (ZKCO).

16 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus cacicus. 27) Male, dorsal view; 28) Male, ventral view; 29) Male, head, dorso-lateral view; 30) Medium sized male, mandibles; 31) Small male, mandibles; 32) Male, protibia; 33) Male, protarsus; 34) Male, metatarsus; 35) Male, mesotarsus; 36) Male, wing; 37) Female, dorsal view; 38) Female, ventral view; 39) Femle, metatarsus.

17 216 THE COLEOPTERISTS BULLETIN 67(3), 2013 Additionally, we examined the following three males: Mexico, male (ex Desbrochers Collection; no other data) (IRSN); 2 males (no other data) (ZMHB). We believe that the place Mexico is only an indication of the type locality of the species. Types and Type Locality. Of Prionacalus cacicus (Figs ): As mentioned above, White (1845) affirmed that there were three specimens from Mexico. Thus, P. cacicus was described from three syntypes. In 1850, White stated that the specimen figured by him as being a female of P. cacicus was also a male, and named it P. iphis. White (1850) did not record the gender of the third specimen and did not provide a figure. He only affirmed that both specimens figured were males and belonged to different species. Waterhouse (1872) stated that: The British Museum has received a pair ( and ) ofan insect from Peru, which I am unable to separate from Pr. cacicus, White, although the sizes of the two males are very different; that from Peru measuring (including the mandibles) 30 lines, whilst the type of Pr. cacicus is only 17 lines. Waterhouse (1900) wrote: The British Museum has recently received a few specimens of Longicorns of the genus Prionacalus. One I refer with a slight doubt to P. cacicus, White, but in the type the tubercle behind the eye is more directed backwards than in the specimen just received. Finally, Lameere (1910) commented on P. iphis(translation): I saw the type in the British Museum; only one male is known. From those three statements, we conclude that only one male of P. cacicus and another of P. iphisare deposited in the BMNH as types of those species. In addition, we conclude that White (1845) either mistakenly recorded three specimens, when actually there were two, or that one of the types has been lost. The website of the BMNH (2013) records that there is only one type of Prinocalus [sic] cacicus deposited there (sex not indicated): holotype. EHN examined the types in the BMNH and found only one specimen labeled as the type of P. cacicus and another as the type of P. iphis. According to the ICZN (1999, Article ): The type series of a nominal species-group taxon consists of all the specimens included by the author in the new nominal taxon And Article 72.6 (ICZN 1999) points out: The fact that a specimen is already the name-bearing type, or part of the name-bearing type, of one nominal species-group taxon does not prevent its being the name-bearing type, or part of the name-bearing type, of another. Thus, P. cacicus does not have a holotype: it has three syntypes. At least one of those specimens is the type of P. iphis, but it is not possible to affirm if this species has only one type (the holotype) or two syntypes. Although White (1850) recorded that The other specimen may be named Prionacalus Iphis, he was only affirming that the specimen figured is P. iphis. In our opinion, it is incorrect to infer that White was affirming that only the specimen figured belonged to this species. According to Recommendation 73F (ICZN 1999): Where no holotype or syntype was fixed for a nominal species-group taxon established before 2000, and when it is possible that the nominal species-group taxon was based on more than one specimen, an author should proceed as though syntypes may exist and, where appropriate, should designate a lectotype rather than assume a holotype. Also, according to Article 74.6 (ICZN 1999): When it has been accepted that a nominal species-group taxon was based on a single specimen and the original description neither implies nor requires that there were syntypes, and if it is considered subsequently that the original description was based on more than one specimen, the first author to have published before 2000 the assumption that the species-group taxon was based upon a single type specimen is deemed to have designated that specimen as the lectotype. Thus, it is possible to affirm that Waterhouse (1872) designated as lectotype of P. iphis the specimen figured as female of P. cacicus in White (1845: figs. 2, 2a, 2b). It is important to note that the lectotype male of P. iphis also remains as a syntype (paralectotype) of P. cacicus. White (1845) indicated that the three specimens were purchased by Mr. Gray for the British Museum from M. Hartweg, and that the specimens were from Mexico. However, there is no record of any species of Prionacalus collected in Mexico, except those by White (1845, 1850, 1851), and of authors who followed White. Thomson (1877) also questioned Mexico as a type locality. According to the ICZN (1999, Recommendation 76A): 76A.1. In ascertaining or clarifying a type locality (and type horizon, type host, and similar terms), an author should take into account: 76A.1.4. as a last resort, and without prejudice to other clarification, localities within the known range of the taxon or from which specimens referred to the taxon had been taken. Prionacalus cacicus is currently recorded in Ecuador and Peru. The species also occurs in Colombia. Thus, the indication of Popayau [Popayán] and Bogotá (Colombia) (Thomson 1877) must be taken into account. Therefore, following Thomson (1877) we indicate the type locality of P. cacicus as Andes (South America). It is not possible to determine a precise locality in this region (following information provided by Thomson), because he mentioned more than one place in the Andes. It is important to note that there are at least two places in Colombia named Mexico located in two different departments: Huila and Arauca.

18 THE COLEOPTERISTS BULLETIN 67(3), The former is in the Andes (2,368 m elevation) and not far from places where other species of Prionacalus are known to occur. As seen in Fig. 108, there is no other information on the type s labels. Of Psalidognathus Wallisi: Described based on a couple of specimens from Ecuador (Loja). As mentioned above, Lameere (1910) affirmed that the male specimen corresponds to P. modestus but the female to P. cacicus. Even so, according to the ICZN (1999: Article ), both are types of P. wallisi. The syntypes are deposited at MLUH. Of Prionocalus [sic] gunteri (Figs ): Holotype male from Ecuador (Zaruma, not Zoruma as originally recorded, Department of El Oro), deposited at BMNH. Remarks. Figure 2, plate 2 in Lameere (1919) clearly does not correspond to P. buckleyi. It distinctly corresponds to P. cacicus. According to Lameere (1910) (translation): P. Gunteri Gahan was based on a male with abdomen finely punctate, elytra separately rounded at apex, appendages reddish, humeri very prominent, elytra less rough towards apex. Lameere (1913, 1919) recorded P. gunteri as a variety of P. buckleyi without explanation. Thus, Lameere (1913) revalidated the former species (not in 1910 as pointed out by Monné 1995, Monné 2006). According to the ICZN (1999: ), on determination of subspecific or infrasubspecific rank of names following a binomen: it is subspecific if first published before 1961 and its author expressly used one of the terms variety or form Since Lameere (1913) considered P. gunterias a variety of P. buckleyi, it is correctly considered a subspecies. It is important to note that Quentin and Villiers (1983) and Hüdepohl (1985) mentioned only P. buckleyi, without a subspecies. Prionacalus gunteri is not a synonym of P. buckleyi, as currently considered, or even a subspecies of the latter. The cephalic carinae, prothoracic shape, and elytral sculpture are as in some specimens of P. cacicus, and not as in P. iphis. Prionacalus demelti Quentin and Villiers, 1983 (Figs. 7, 13, 40 50, 88 91) Prionocalus demelti Quentin and Villiers 1983: 224, 2 figs.; Hüdepohl 1985: 122 (key); Pardo- Locarno 2006: 234. Prionacalus demelti; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 83 (catalogue); Martínez 2000: 85 (distribution); Komiya 2001: 31 (figs. 6 8), 32 (key); Di Iorio 2003: 3 (distribution); Salazar 2005: 246; Arai 2005: 34, fig. 2 (male); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 16 (males, females), 17 (male), 110 (female). Redescription. Male (Fig. 40). Integument black to dark brown, usually dorsally darker; maxillary and labial palpi reddish brown; femora from brown to reddish brown, always apically blackish; tibiae from brown to reddish brown, basally blackish or dark brown; tarsi reddish brown (at least, distal 1/3 of claws always black); scape blackish to dark brown; antennomeres gradually lighter from III to V, and VI-XI with same color as V; elytra with irregular and variable reddish areas (sometimes almost absent). Head: Coarsely punctate-rugose, mainly between cephalic carinae on region of eyes and antennal tubercles; area between cephalic carinae distinctly depressed, centrally with distinct sulcus from clypeus to area of posterior ocular edge (sometimes reaching only middle of eyes). Antennal tubercle coarsely confluently punctate on base, almost smooth near apex; bases far from each other (distance between them equal to about 1.2 times tubercle width). Cephalic carinae (Fig. 42) convergent from antennal tubercle to after eyes, divergent from this point to apex (sometimes subparallel), strongly elevated at apex, forming spine-like tops. Area behind lower ocular lobes with abundant small asperities. Lateral tubercle of head very large, acute at apex. Subhorizontal area of clypeus coarsely, shallowly or smoothly punctate, usually with distal margin elevated. Distal area of clypeus distinctly concave, smooth. Labrum subtriangular, apex acute or slightly rounded. Distance between upper ocular lobes times longest width of 1 lobe. Gena (Fig. 42) with long projection directed forwards (sometimes downward). Submentum depressed, coarsely, moderately abundantly punctate. Mandible (Fig. 43) as long as head to slightly longer; subcylindrical on base, distinctly laminar towards apex; surface coarsely, abundantly punctate, gradually finer, sparser towards apex (smooth close to inner margin); both mandibles with inner margin pluridentate from base to distal plate, the latter also with numerous, irregular small teeth at its basal 1/2. Longest width of last maxillary (Fig. 7) and labial palpomeres equal to about 0.25 times length. Antennae reaching or slightly surpassing elytral apex; scape subcylindrical, coarsely, abundantly, confluently punctate on basal 1/2, sparsely, finely punctate towards apex; antennomere III about 1.6 longer than scape; antennomeres III-X (Fig. 13) with projection on both sides of apex, notably long on III-V. Thorax: Latero-anterior angles of prothorax projected forward, almost acute; latero-posterior angles of prothorax not projected sideward, rounded; lateral margin with 3 distinct teeth, the 2 anterior teeth protracted, the most anterior smaller. Pronotum with distinct depression on center of disc, sometimes longitudinally divided by slightly distinct

19 218 THE COLEOPTERISTS BULLETIN 67(3), 2013 carina; coarsely, abundantly punctate (at least partially confluent laterally). Prosternal process with rounded projection at underside of apex (sometimes slightly distinct). Elytra coarsely vermiculate throughout; apex variable: individually rounded or rounded together; with or without projection at sutural angle (occasionally, left elytron shorter than right). Apical tooth humeral projection large. Abdomen: Usually not surpassing elytral apex (sometimes slightly surpassing). Ventrites I-IV (Fig. 41) very finely, sparsely punctate; ventrite V coarsely, abundantly punctate. Legs: Metafemora surpass elytral apex by about middle. Ventral sulcus of protibiae visible only between about middle and before apex. Protarsi (Fig. 44) moderately elongate and narrow; protarsomeres I-III with denticle at each apex. Mesotarsi (Fig. 45) slender; apex of mesotarsomeres I-III with distinct spine at each apex. Metatarsi (Fig. 46) distinctly slender; apex of metatarsomeres I-III spiny. Female (Fig. 47). Dorsally blackish, ventrally dark brown; distal area of ventrites blackish; scape brownish; antennomeres reddish brown. Mandibles (Fig. 49) laminar; inner margins pluridentate. Antennae reaching distal 1/3 of elytra. Longest width of last maxillary and labial palpomeres about 0.20 times length. Cephalic carinae as in males. Distance between upper ocular lobes 3.0 times longest width of 1 lobe. Lateral tubercle of head as in male. Prothorax as in males. Abdomen surpassing or not elytral apex. Ventrites I-IV almost impunctate (Fig. 48); ventrite V very finely, sparsely punctate. Metatarsi (Fig. 50) shorter than in male. Dimensions. Male/female. Total length (including mandibles) = / mm; prothoracic length = / mm; prothoracic width between apices of anterior angles = / mm; prothoracic width between apices of posterior angles = / mm; humeral width = / mm; elytral length = / mm. Geographical Distribution. Colombia. Material Examined. COLOMBIA: Female, XII.1991, [no collector indicated] (ZKCO); male, VII.1992, [no collector indicate] (ZKCO). Huila: Gigante, 1 male, 1 female, 1974, (no collector indicated) (ZKCO); 1 male, 1976, (no collector indicated) (ZKCO), 1 male, 1 female, [no date indicated], local collector (MPCO); male, XI.1978, [no collector indicated] (DHPC). Boyacá: near Muzo, male, V.1983, [no collector indicated] (ZKCO). Yari: Caqueta, 2 males, I.1982, (no collector indicated) (ZKCO). Types and Type Locality. Prionocalus [sic] demelti was described based on a holotype male (Figs. 88, 89) and an allotype female (Figs. 90, 91), both from Colombia (Putumayo River Valley), deposited at MNHN. Additionally, there were four paratypes (three males, one female) deposited in Carl von Demelt Collection (currently deposited at SMNS). Remarks. Prionacalus demelti resembles Psalidognathus superbus Fries, 1833 by its antennomeres which are distinctly spiny at apex, and by its long genal projection. Those features make this species one of the more easily recognizable Prionacalus species. Prionacalus whymperi Bates, 1892 (Figs. 6, 14, 51 63, 92 95, ) Prionocalus Whymperi Bates 1892: 36, 1 fig; Campos, 1921: 87, fig. 16. Prionocalus whymperi; Quentin and Villiers 1983: 223 (revalidation, lectotype). Prionacalus whymperi; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 61 (catalogue), 2006: 84 (catalogue); Komiya 2001: 31, 32 (key); Di Iorio 2003: 3 (distribution). Psalidognathus (Prionocalus) Buckleyi; Lameere 1910: 376 (part), 1913: 65 (catalogue, part), 1919: 121, pl. 6, fig. 2 (part). Prionocalus giovannii Hüdepohl 1985: 117, 4 figs. New synonymy. Prionacalus giovannii; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 83 (catalogue); Komiya 2001: 32 (key); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist). Prionacalus giovanni; Di Iorio 2003: 3 (distribution). Prionocalus trigonodes Bates 1892: 37; Whymper 1892: 10; Kolbe 1902: 481; Campos 1921: 87; Quentin and Villiers 1983: 224 (lectotype, synonymy); Hüdepohl 1985: 121. New synonymy. Psalidognathus (Prionocalus) trigonodes; Lameere 1910: 377. Redescription. Male (Fig. 51). Head, mandibles, and thorax blackish (sometimes prosternal process reddish brown); maxillary and labial palpi brown to reddish (usually slightly darker on basal segments); scape and pedicel blackish; antennomere III entirely blackish to entirely dark brown (sometimes dark brown only distally); antennomeres IV-XI dark brown (sometimes slightly reddish); elytra dark brown on basal 1/3, gradually reddish towards apex; ventrites dark brown, with distal margin blackish (sometimes, ventrite V almost entirely reddish, except on distal margin); profemora dark brown; mesoand metafemora reddish on basal 1/3, dark brown on remaining surface (occasionally almost entirely dark brown); protibiae dark brown on basal 1/3, gradually reddish towards apex (sometimes reddish

20 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus demelti. 40) Male, dorsal view; 41) Male, ventral view; 42) Male, head, dorso-lateral view; 43) Male, mandibles; 44) Male, protarsus; 45) Male, mesotarsus; 46) Male, metatarsus; 47) Female, dorsal view; 48) Female, ventral view; 49) Female, mandibles; 50) Female, metatarsus.

21 220 THE COLEOPTERISTS BULLETIN 67(3), 2013 only on distal 1/3); meso- and metatibiae brown on base, slightly reddish towards apex (occasionally entirely dark brown); tarsi entirely dark brown to entirely reddish (sometimes dark brown with some parts reddish), except claws that are blackish. Head: Coarsely scabrous between apex of cephalic carinae and prothorax (sometimes only punctate near prothorax), vermiculate between cephalic carinae, coarsely punctatevermiculate between upper ocular lobes and prothorax; area between cephalic carinae distinctly depressed, centrally with distinct sulcus from clypeus to area after antennal tubercles (sometimes from clypeus to almost apex of cephalic carinae); distal margin of vertex (close to abrupt declivity) centrally distinctly emarginate. Antennal tubercle coarsely punctate-vermiculate on base, gradually finer towards impunctate area of apex; bases near each other (distance between them about 0.15 times width or slightly narrower). Cephalic carinae (Fig. 53) distinct, elevated from base to apex (usually less so between antennal tubercle and base of eye); convergent from antennal tubercle to little after eyes, subparallel towards apex; apex not or slightly elevated, not forming a triangular tubercle, but usually wider at this area. Area behind lower ocular lobes variable: not finely striate, and with abundant small asperities; finely striate near prothorax, and with abundant asperities near eyes. Lateral tubercle of head small to moderately large, rounded at apex (Fig. 96) to distinctly acute (Fig. 51). Subhorizontal area of clypeus finely, coarsely, abundantly punctate (occasionally with 1 tubercle on central area). Distal area of clypeus distinctly concave, entirely smooth to slightly rugose at base. Labrum distinctly concave, usually with central longitudinal sulcus, and apex centrally emarginate. Distance between upper ocular lobes times longest width of 1 lobe. Gena rounded at apex (Fig. 53), without downward projection. Submentum depressed, laterally confluently, coarsely punctate, remaining surface coarsely vermiculate, or abundantly punctate, or almost smooth. Mandibles (Fig. 54) times head length; inner side laminar; surface coarsely, abundantly punctate, gradually finer towards apex (smooth near inner margin); left mandible with 1 distinct tooth on basal 1/2 (sometimes another small one nearer base); right mandible with 2 jointly protracted teeth near base (posterior tooth smallest), and another large tooth close to the base of distal plate; distal plate of left mandible wider than that of right mandible. Longest width of last maxillary (Fig. 6) and labial palpomeres almost equal to 0.4 times length. Antennae not or slightly surpassing elytral apex; scape slightly enlarged from base to apex, coarsely, confluently punctate; antennomere III usually distinctly dentate on inner side of apex, rounded or slightly dentate on outer margin, about 1.6 times longer than scape; antennomere IV dentate on both sides of apex (sometimes only on outer side); antennomeres V-X usually slightly projected on both sides of apex. Thorax: Latero-anterior angles of prothorax slightly projected forward, rounded (occasionally somewhat acute); latero-posterior angles of prothorax distinctly projected sideward (tooth-like), acute to moderately rounded at apex; lateral margin with 3 distinct teeth (anterior one distinctly smaller to subequal to middle tooth). Pronotum with a distinct, transverse depression on distal 1/2; centrally with longitudinal carina from near base to apex; coarsely punctate-vermiculate, usually with smooth, narrow band on base; anterior margin distinctly emarginate centrally. Prosternal process with spiniform projection at underside of apex. Elytra coarsely vermiculate throughout; apices individually to jointly rounded (sometimes moderately truncate). Apical tooth of humeral projection small to very large. Abdomen: Not surpassing elytral apex. Ventrites (Fig. 52) moderately coarsely, abundantly punctate (mainly I-IV laterally and V throughout), or distinctly sparsely punctate on ventrites I-V (mainly centrally) and V abundantly punctate. Legs: Apex of metafemora surpasses elytral apex by about 0.2 times length of metafemora. Ventral sulcus of protibiae distinct only from middle to near apex (usually present after basal 1/3, but not distinctly marked); protibiae enlarged near middle (Fig. 58). Protarsi (Fig. 55) moderately short and wide; protarsomeres I-III with small denticle at apex. Mesotarsi (Fig. 56) slender; apex of mesotarsomeres I-III with distinct denticle at apex. Metatarsi (Fig. 57) distinctly slender; apex of metatarsomeres I-III spiny (mainly III). Female (Fig. 59). Head and prothorax dark brown, with some parts blackish; mandibles blackish; abdomen entirely dark brown or reddish brown with apex blackish; scape dark brown; pedicel and antennomeres III-XI reddish brown. Inner margin of mandibles (Fig. 61) laminar; inner margins similar to those in males but with only 1 tooth between base and distal plate. Antennae slightly surpassing middle of elytra. Longest width of last maxillary and labial palpomeres about 0.35 times length. Cephalic carinae as in males, or distinctly divergent after eyes. Distance between upper ocular lobes times longest width of 1 lobe. Lateral tubercle of head as in male, with the same variations. Prothorax as in males (in 2 females examined, anterior lateral tooth smaller than middle tooth). Prosternal process as in males (Fig. 14). Abdomen distinctly surpassing elytral apex. All ventrites (Fig. 60) moderately coarsely, sparsely punctate. Mesoand metatarsi shorter than in male; tarsomeres

22 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus whymperi. 51) Male, dorsal view; 52) Male, ventral view; 53) Male, head, dorso-lateral view; 54) Male, mandibles; 55) Male, protarsus; 56) Male, mesotarsus; 57) Male, metatarsus; 58) Male, protibia; 59) Female, dorsal view; 60) Female, ventral view; 61) Female, mandibles; 62) Female, narrow metatarsus; 63) Female, thick metatarsus.

23 222 THE COLEOPTERISTS BULLETIN 67(3), 2013 slender and elongate (Fig. 62) or thick and short (Fig. 63). Dimensions. Male/female. Total length (including mandibles) = / mm; prothoracic length = / mm; prothoracic width between apices of anterior angles = / mm; prothoracic width between apices of posterior angles = / mm; humeral width = / mm; elytral length = / mm. Geographical Distribution. Ecuador. Material Examined. ECUADOR: 2 males, 2 females, [no other data], Fritsche V. col. (ZMHB); male, VI.1982, J. P. Marechal col. (ZKCO). Cotopaxi: 2500m,1male,1female, III.1985, P. Arnaud col. (MPCO). Pichincha: female, IV.2001, [no collector indicated] (ZKCO); 2 males, 1 female, XII.2001, [no collector indicated]; Los Alpes (2000 m), male, 12.VII.1935, Schultze- Rhonhof S. G. col. (ZMHB); Santo Domingo de los Colorados, male, VII.2001, [no collector indicated] (ZKCO). Types and Type Localities. Of Prionacalus whymperi (Figs ): Described from two males and one female, all from Milligalli (6200 feet). According to Myers (1969): from Milligalli, Ecuador, a locality at about 1900 meters elevation on the western slopes of Cerro Corazon (Andrade Marin, 1931, p. 30). The type locality is located in the province of Pichincha. The syntypes are deposited at MNHN. Quentin and Villiers (1983) stated (translation): Among the three syntypes, we designate a LECTOTYPE male with 38 mm length (without mandibles) and with the following labels: Milligalli, Ecuador feet. Ed. Whymper., Ex Musaeo H. W. Bates, 1892, Museum Paris, Coll. H. W. Bates, 1952, Whymperi Bates. LECTOTYPE female with 63 mm length (without mandibles) with the same labels and: Prionocalus Whymperi Bates,. One PARALECTOTYPE male from same place. According to the ICZN in effect at that time (ICZN 1964) and currently (ICZN 1999), those designations are invalid, because a species can only have one lectotype: Designation of a lectotype. A lectotype may be designated from syntypes to become the unique bearer of the name of a nominal speciesgroup taxon and the standard for its application; and Lectotype designations before In a lectotype designation made before 2000, either the term lectotype, or an exact translation or equivalent expression (e.g. the type ), must have been used or the author must have unambiguously selected a particular syntype to act as the unique name-bearing type of the taxon. We designate here as the lectotype for P.whymperi the specimen (Figs ) designated by Quentin and Villiers (1983) as the lectotype male. The specimen has the following labels (Fig. 99): 1. White [Handwritten]: whymperi / Bates; 2. White: Milligalli [Handwritten] / Ecuador. (Printed) / 6000 [Handwritten] feet [Printed] / Ed. Whymper [Printed]; 3. White [Printed]: Ex-Musaeo H. W. Bates / 1892; 4. White [Printed]: Museum Paris / Coll. H. W. Bates / White: Prionocalus [sic] whimperi[sic] Bates [Handwritten] / Lectotype [Handwritten] / Quentin and Villiers det. 19 [Printed] 82 [Handwritten]; 6. Red [Printed]: Lectotype 7. Red and yellow [Printed; added by us]: LECTOTYPE / Prionacalus whymperi Although Bates (1892) recorded 6200 feet, the type label indicates the elevation as either 600 or 6,000 ft. We can never know which is the true value recorded since the pin was inserted exactly where the last zero would be placed. Of Prionocalus [sic] giovannii: Holotype and paratype males, from Ecuador (Province of Pichincha, Santo Domingo de los Colorados), deposited at ZSMC. Of Prionocalus [sic] trigonodes (Figs ): The number of specimens originally included in the description is controversial. Bates (1892) recorded: Hab. La Mona (under 200 feet), one. Taken also by Mr. Buckley on his last journey. However, he also indicated: Long milim.. Quentin and Villiers (1983) designated the lectotype for the species (translation): We designated as LECTOTYPE a male with 37 mm long (without mandibles), with the following labels: Ecuador, feet. Ed. Whymper, La Mona, Ex Museum H. W. Bates, trigonodes Bates, Museum of Paris, Coll. H. W. Bates. La Mona is located in the province of Los Rios, Ecuador. The species was likely described based on a single specimen. According to Whymper (1892): Amongst the few species secured on the first day s journey, there have been found an undescribed Ant (Camponotus), a Bug (Pnohirmus), and two Beetles (Epitragus and Prionocalus [sic]). These are described and figured in the Supplementary Appendix with is published simultaneously with this volume. The Prionocalus [sic] that is described by Mr. H. W. Bates under the name P. trigonodes was picked up close to La Mona. Based on this information, we believe that Bates had only a single specimen when he described the species (indicating it was the holotype). Thus, the designation of a lectotype by Quentin and Villiers (1983) was in error, based on the incorrect information provided by Bates (1892).

24 THE COLEOPTERISTS BULLETIN 67(3), Remarks. As mentioned above, Lameere (1910) considered P. whymperi as a synonym of P. buckleyi (translation): I have not seen the type of P. Whymperi Bates; the species is established on a male slightly differing from P. Buckleyi, according to Bates himself, but also on a female with elytra notably surpassing elytral apex. I have before me a male of P. Buckleyi that also has this feature, but I cannot see more than an individual variation. Quentin and Villiers (1983) revalidated the species (translation): The study of the types of Bates, preserved in the Paris Museum, showed us that Lameere (loc. cit.), who had not seen them, committed various mistakes that we rectify as follows [translation]: Lameere (loc. cit.: 731) had arbitrarily placed this species in synonymy of P. buckleyi Waterhouse. Instead, we consider it as valid [NEW COMBINATION]. They are distinguished by the following characters: P. whymperi Bates: 1. Anterior tibiae of male 6 times longer than wide, dilated at middle, and then narrowed at apical fourth; setae of inner face starting after basal fourth; 2. Tarsi slender in both sexes, second mesotarsomere about as long as wide at its apex; 3. Elytra covering the abdomen. P. buckleyi Waterhouse: 1. Anterior tibiae of male 7 times longer than wide, enlarged from base to the apex; setae of inner face extending on entire length, from the base; 2. Tarsi thicker; second mesotarsomere approximately one and a half time longer than wide at its apex; 3. Elytra usually leaving visible the apex of abdomen. We agree with Quentin and Villiers (1983). Prionacalus whymperi is not a synonym of P. buckleyi, and thus, not a synonym of P. i p h i s. The main character separating P. whymperifrom P. iphisis the shape of the sulcus on the inferior side of the protibiae, starting far from the base in the former and near the base in the latter. Although Hüdepohl (1985) was aware of the work by Quentin and Villiers (1983), he excluded P. whymperiin his key to the species of Prionocalus [sic] without explanation. Prionacalus whymperi was listed as a valid species in Monné (1995, 2006). However, it was considered a synonym of P. buckleyi in Monné and Giesbert (1994), Monné and Hovore (2005, 2006), and it was omitted in Jeniš (2010). Based on the specimens examined, original descriptions, and photographs of the types, we conclude that P. giovannii is a junior synonym of P. w h y m p e r i. It is strange that Hüdepohl did not observe the information from Quentin and Villiers (1983) on the protibiae of P. whymperi: dilated at middle setae of inner face starting after basal 1/4. This information would have shown that the protibiae in the types of P. w h y m p e r i are exactly as in the holotype of P. giovannii. This feature and the shape of the first spine on the lateral margin of the prothorax can be used to distinguish P. w h y m p e r i from the other species of the genus. As mentioned above, Quentin and Villiers (1983) considered P. trigonodes another synonym of P. buckleyi (translation): Lameere (loc. cit., Revision: 732), without seeing the type, assigned to this species erroneous characters based on specimens wrongly identified in the British Museum. In fact, trigonodes corresponds to small individuals of buckleyi and the name cannot be maintained. The short description of P. trigonodes does not allow it to be differentiated from P. buckleyi.however, examining the detailed photographs of the lectotype male, we conclude that it is equal to P. w h y m p e r i and not to P. buckleyi (= P. iphis). Although the protibiae (Fig. 92) are not distinctly enlarged from the middle, they are somewhat slender, and the ventral sulcus does not start very near the base. In addition, the prosternal process has a distinct, spiniform projection on the underside of the apex, a feature that was not observed in any specimen of P. iphis. Also, although Quentin and Villiers (1983) had affirmed that P. trigonodes agrees with small specimens of P. buckleyi, the length of the holotype (their lectotype), 37 mm, is about as long as the lectotype of P. buckleyi (18 lines). One English line corresponds to about 2.12 mm ( mm). Thus, the lectotype male of P. buckleyi is about 38.2 mm in length. Both P. whymperi and P. trigonodes were recorded as having been described by Bates in However, as mentioned above, Whimper (1892) affirmed: in the Supplementary Appendix which is published simultaneously with this volume. Thus, it is possible to infer that the date on the Supplementary Appendix was pre-dated. In other words, it was printed first in 1891 but not distributed until Prionacalus woytkowskii (Heyrovsky, 1960) (Figs. 3, 9, 10, 64 75) Psalidognathus (Prionocalus) woytkowskii Heyrovsky 1960: 162, 3 figs. Prionocalus woytkowskii; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121. Prionacalus woytkowskii; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 61 (catalogue), 2006: 84 (catalogue); Komiya 2001: 29, 30 (figs. 2, 3), 32 (key); Di Iorio 2003: 3 (distribution); Arai 2005: 34, fig. 3 (male), 4 (female); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 16 (males, females), 17 (male), 124 (male), 125, female; Monné et al. 2012: 7 (checklist).

25 224 THE COLEOPTERISTS BULLETIN 67(3), 2013 Prionacalus buckleyi; Jeniš 2001: 27 (fig. 91/55). Prionacalus giovannii; Jeniš 2010: 19 (male, female), 114 (male), 115 (female); Monné et al. 2012: 7 (checklist). Redescription. Male (Fig. 64). Integument black (sometimes with small dark brown areas); maxillary and labial palpi reddish brown; tibiae reddish on distal fourth (sometimes almost from basal 1/3); tarsi from brown to reddish brown, except small dark area at apex and blackish claws; pedicel and antennomere III blackish to dark brown, except distal 1/6 of antennomere III reddish; basal 3/4 of antennomere IV brown or dark brown, distal 1/4 from brown to reddish; antennomere V brown to reddish; antennomeres VI-XI reddish. Head: Coarsely punctate-vermiculate, mainly between cephalic carinae; area between cephalic carinae distinctly depressed, centrally with distinct sulcus from clypeus to area of posterior ocular edge. Antennal tubercle coarsely punctatevermiculate on base, becoming smoother towards apex (only with very sparse coarse punctures); bases moderately far from each other (distance between them equal to almost each width). Cephalic carinae (Fig. 66) slightly distinct by the coarse sculpture of dorsal surface of head but usually more or less recognizable; subparallel from base to apex. Area behind lower ocular lobes with abundant small asperities. Lateral tubercle of head large, rounded at apex (Fig. 64). Subhorizontal area of clypeus smooth, usually centrally with 2 small callosities. Distal area of clypeus distinctly concave, smooth. Labrum distinctly concave (occasionally slightly concave), apex centrally emarginate or not. Distance between upper ocular lobes times longest width of 1 lobe. Gena acute at apex (Fig. 66), with short sideward projection (sometimes moderately truncate and not sideward projected). Submentum depressed, transversely sulcate (sometimes vermiculate), finely to moderately coarsely punctate (punctures moderately sparse; occasionally absent). Mandibles (Fig. 67) times head length; inner margin subcylindrical at basal 1/3; surface coarsely, abundantly punctate on basal 1/3, gradually finer, sparser towards apex (smooth near inner margin); left mandible with 1 distinct tooth near base of distal plate; right mandible with 1 large tooth close to base of distal plate (in small specimens, usually with another smaller tooth jointly protracted); distal plate of left mandible wider than that of right mandible. Longest width of last maxillary (Fig. 3) and labial palpomeres about 0.45 times length. Antennae slightly to distinctly surpassing elytral apex; antennomeres moderately slender (Fig. 10) to wide (Fig. 9); scape slightly to distinctly enlarged from base to apex, coarsely, confluently punctate, except on distal 1/5 which is almost smooth; antennomere III rounded on both sides of apex, about 1.4 times longer than scape; antennomeres IV-X dentate on outer apex, rounded to angulate on inner apex. Thorax: Lateroanterior angles of prothorax slightly projected forward, rounded; latero-posterior angles of prothorax distinctly projected sideward (tooth-like), moderately rounded at apex; lateral margin with 3 distinct teeth (anterior tooth smallest). Pronotum with distinct depression on center of disc (occasionally slightly marked); coarsely punctatevermiculate (laterally less so), occasionally with small, impunctate, shiny callosity on each side of anterior third; anterior margin distinctly emarginate medially. Elytra very coarsely vermiculate throughout; apex dentate (occasionally not) at sutural angle (frequently, left elytron shorter than right one). Apical tooth of humeral projection small to very large. Abdomen: Not surpassing elytral apex. Ventrites I-IV finely, sparsely punctate (laterally slightly more abundantly) (Fig. 65); ventrite V abundantly, moderately coarsely punctate. Legs: Apex of metafemora surpassing elytral apex by about 1/2 length of metafemora (occasionally, about 0.4 times). Ventral sulcus of protibiae distinct from near base to near apex. Protarsi (Fig. 68) moderately short and wide; protarsomeres I-III with small denticle at apex. Mesotarsi (Fig. 69) slender; mesotarsomeres I-III with distinct, apical denticle (mainly on III). Metatarsi moderately (Fig. 71) to distinctly (Fig. 70) long and slender; apex of metatarsomeres I-III spiny. Female (Fig. 72). Integument dark brown; mandibles blackish; scape brown; antennomeres III-XI reddish; elytra with or without some irregular reddish areas; color of palpi and legs as in males. Inner margin of mandibles (Fig. 74) laminar; inner margin of left mandible as in male, but with tooth wider; inner margin of right mandible with 2 teeth jointly protracted, close to the distal plate. Antennae slightly surpassing middle of elytra (sometimes almost reaching distal 1/3). Longest width of last maxillary and labial palpomeres about 0.3 times length. Cephalic carinae as in males. Distance between upper ocular lobes equal to about 3.0 times longest width of 1 lobe. Lateral tubercle of head as in male. Prothorax as in males. Abdomen distinctly surpassing elytral apex. All ventrites (Fig. 73) finely, sparsely punctate. Meso- and metatarsi (Fig. 75) shorter than in male. Dimensions. Male/female. Total length (including mandibles) = / mm; prothoracic length = / mm; prothoracic width between apices of anterior angles = / mm; prothoracic width between apices

26 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus woytkowskii. 64) Male, dorsal view; 65) Male, ventral view; 66) Male, head, dorso-lateral view; 67) Male, mandibles; 68) Male, protarsus; 69) Male, mesotarsus; 70) Male, narrow metatarsus; 71) Male, thick metatarsus; 72) Female, dorsal view; 73) Female, ventral view; 74) Female, mandibles; 75) Female, metatarsus.

27 226 THE COLEOPTERISTS BULLETIN 67(3), 2013 of posterior angles = / mm; humeral width = / mm; elytral length = / mm. Geographical Distribution. Peru (Heyrovsky 1960), Bolivia (Monné 2006), and Argentina (Di Iorio 2003). Material Examined. PERU: Acomayo, male, III.1984 [no collector indicated] (ZMHB). Ucayali: Pucallpa, male, 12.XII.1984, [no collector indicated] (DDPC); female, 15.XII.1984, [no collector indicated] (DDPC). Huanuco: male, IX.1984, local collector (MPCO); Cordillera of Carpish, male, 01.X.1950, [name of collector unreadable] (EMEC); female, II.1986, [no collector indicated] (DDPC); (2,800 m), 2 males, II.2009, local collector (MPCO); male, local collector (MPCO); (Carpish Pass; 2600 m), 3 males, III.1983, [no collector indicated] (DHPC); Tingo María, 1 male, 1 female, XII.2003, local collector (MPCO); (680 m), 1 male, 1 female, 04.V.1973, [no collector indicated] (SNSD), 1 male 1 female, 10.VI.1974, (no collector indicated) (ZKCO). Junín: Satipo, female, VII.1991 [no collector indicated] (DDPC), 5 males, I.1990 (no collector indicated) (ZKCO). Pasco: Oxapampa, male, [data unreadable] (IRSN). San Martín: Uchiza (Huallaga River), 20 males, 2 females, I.1985, (no collector indicated) (ZKCO). Types and Type Locality. Holotype, allotype, and two paratypes from Peru (Mount Carpish, 2,813 m, between Huánuco and Tingo María, Andes). Heyrovsky did not record the sex of the types. The holotype appears to be a male. The types are deposited in the Leo Heyrovsky Collection (currently at NMPC). Remarks. Hüdepohl (1985) stated that P. woytkowskii was a likely a synonym of P. trigonodes. However, he did not comment on the affirmation by Quentin and Villiers (1983), who said that P. trigonodes is a synonym of P. buckleyi. Also, Komiya (2001) considered P. woytkowskii and P. trigonodes as synonyms, following Hüdepohl (1985). Hüdepohl (1985) and Komiya (2001) included those species in their keys, following the couplet head without clear cephalic carinae. However, the lectotype male of P. trigonodes has distinct cephalic carinae. We examined an aberrant specimen with elytra not entirely vermiculate, but agreeing in all other characters with the types. Prionacalus iphis White, 1850 (Figs. 4, 5, 15, 76 87, ) Prionacalus Iphis White 1850: 11, 1851: 71; Gemminger and Harold 1872: 2755 (catalogue). Prionacalus iphis; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 83 (catalogue); Komiya 2001: 30 (fig. 5), 32 (key); Di Iorio 2003: 3 (distribution); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 16 (holotype male), 123 (male). Prionocalus Iphis; Lacordaire 1868: 42 (note); Waterhouse 1872: 261 (key); Thomson 1877: 260. Psalidognathus (Prionocalus) Iphis; Lameere 1910: 375, 1913: 65 (catalogue), 1919: 121. Psalidognathus (Prionocalus) iphis; Blackwelder 1946: 555 (checklist). Prionocalus iphis; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121 (key). Prionus (Prionacalus) Cacicus White 1845: 110, pl. 8, fig. 2 (female, not male), 1850: 11 (synonymy). Prionocalus Buckleyi Waterhouse 1872: 261; Thomson 1877: 260; Whymper 1892: 10; Nonfried 1892: 17; Lameere 1883: 6 (catalogue); Bates 1892: 37; Wood 1892: 233, fig. 10; Campos 1921: 87. New synonymy. Prionacalus Buckleyi; Waterhouse 1880: 485. Prionacalus buckleyi; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 60 (catalogue), 2006: 82 (catalogue); Komiya 2001: 32 (key); Di Iorio 2003: 3 (distribution); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 19 (syntypes male and female), 116 (male), 117 (female). Psalidognathus (Prionocalus) Buckleyi; Lameere 1910: 376 (part), 1913: 65 (catalogue, part), 1919: 121 (part). Psalidognathus (Prionocalus) buckleyi; Blackwelder 1946: 555 (checklist). Prionocalus buckleyi; Quentin and Villiers 1983: 224; Hüdepohl 1985: 121. Prionocalus Whitei Waterhouse 1900: 505, 1 fig.; Lameere 1910: 376 (synonymy). Psalidognathus (Prionocalus) Buckelyi var. Whitei; Lameere 1910: 65 (revalidation). Prionacalus Emmae Kolbe 1902: 480, pl. 7, figs. 4, 4a; Lameere 1910: 376 (synonymy). Redescription. Male (Fig. 76). Integument black; maxillary and labial palpi brown (somewhat dark) to reddish brown; femora black to dark brown; tibiae entirely black, or black on base and gradually reddish brown towards apex, or entirely reddish brown; tarsi dark brown to reddish brown (at least distal 1/3 of claws always black); antennae entirely black, or scape black and remaining antennomeres brown, or black with distal portion of some antennomeres brownish; distal 1/3 of elytra frequently dark brown (sometimes lighter), but the distal 1/2 sometimes distinctly reddish. Head: Coarsely punctate-rugose, mainly between cephalic carinae on region of eyes and antennal tubercles; area between cephalic

28 THE COLEOPTERISTS BULLETIN 67(3), carinae distinctly depressed, centrally with distinct sulcus from clypeus to area of middle of eyes. Antennal tubercle coarsely, confluently punctate on base, almost smooth towards apex; bases far from each other (distance between them equal to about 1.4 times width). Cephalic carinae (Fig. 78) distinctly convergent from antennal tubercle to after eyes, divergent at its distal 1/3, usually moderately elevated at apex, becoming tuberclelike, with a transverse carina connecting the cephalic carinae near apex (always distinctly lower and narrower than the cephalic carina; frequently interrupted at middle; sometimes almost absent). Area behind lower ocular lobes with abundant, small asperities. Lateral tubercle of head large to moderately small, rounded to almost acute at apex, narrow to wide. Subhorizontal area of clypeus coarsely, shallowly punctate to almost smooth, usually with distal margin elevated. Distal area of clypeus distinctly concave, smooth. Labrum distinctly concave, apex centrally emarginate or not. Distance between upper ocular lobes times longest width of 1 lobe. Gena acute at apex (Fig. 78), downwardly projected (sometimes rounded, not projected). Submentum depressed, coarsely, shallowly, confluently punctate (sometimes moderately coarsely, transversely vermiculate or distinctly finer punctate). Mandibles (Fig. 79) times head length; subcylindrical on basal 1/2 (large males), or more distinctly laminar (small males); surface coarsely, abundantly punctate on basal 1/3, gradually finer, sparser towards apex (smooth closer to inner margin); left mandible in large males with small denticulation between base and distal plate, with a moderately large tooth close to distal plate, rounded at apex, or with distinct teeth between base and distal plate, variable in number, position, and size; left mandible in medium sized and small males with 3 4 distinct teeth (usually part of them jointly protracted) between base and distal plate; inner margin of right of large, medium, and small males as left mandible, but occasionally only with a large, apically rounded tooth or with a large, triangular tooth close to distal plate; distal plate of left mandible wider than that of right mandible. Longest width of last maxillary and labial palpomeres 0.35 times (Fig. 4) to 0.55 times (Fig. 5) length. Antennae reaching only distal 1/5 of elytra, or distinctly surpassing elytral apex; scape slightly enlarged from base to apex (sometimes more distinctly enlarged), coarsely, confluently punctate, except on distal 1/5, which is very sparsely, finely punctate; antennomere III times longer than scape; antennomeres III-X with small projection on both sides of apex (sometimes with distinct spines). Thorax: Lateroanterior angles of prothorax slightly projected forward, rounded; latero-posterior angles of prothorax distinctly projected sideward (toothlike), acute at apex; lateral margin with 3 distinct teeth (anterior tooth smallest). Pronotum with distinct depression on center of disc, longitudinally divided by carina; coarsely punctate-vermiculate (less so laterally), sometimes with distinct, impunctate, shiny area on each side of base; anterior margin centrally emarginate or not. Prosternal process (Fig. 15) without spiniform projection on underside of apex. Elytra coarsely vermiculate throughout; apices individually or jointly rounded; with or without projection at sutural angle (occasionally, left elytron shorter than right one). Apical tooth of humeral projection very small to large. Abdomen: Not surpassing elytral apex. Ventrite I (Fig. 77) moderately finely, sparsely punctate; ventrites II-IV coarsely, abundantly punctate, except near distal margin which is smooth (occasionally, II-IV moderately finely and sparsely punctate, but always denser and coarser than on I); ventrite V abundantly, coarsely punctate throughout. Legs: Apex of metafemora surpassing elytral apex by about 1/5 (occasionally about 2/5) length. Ventral sulcus of protibiae (Fig. 83) distinct from near base to near apex. Protarsi (Fig. 80) moderately short and wide; protarsomeres I-III with small, apical denticle. Mesotarsi (Fig. 81) slender; apex of mesotarsomeres I-III with distinct spine at apex. Metatarsi (Fig. 82) distinctly slender; apex of metatarsomeres I-III spiny. Female (Fig. 84). Head and prothorax blackish brown; mandibles blackish; abdomen entirely dark brown; antennae blackish (pedicel partially reddish). Inner margin of mandibles (Fig. 86) laminar; inner margin similar to those in males, but with only 1 tooth between base and distal plate. Antennae reaching distal 1/3 of elytra. Longest width of last maxillary and labial palpomeres about 0.25 times length. Cephalic carinae as in males. Distance between upper ocular lobes equal to 2.5 times longest width of 1 lobe. Lateral tubercle of head somewhat small, narrow. Prothorax as in males. Abdomen distinctly surpassing elytral apex. Ventrite I (Fig. 85) finely, sparsely punctate; ventrites II-V coarser, more distinctly, abundantly punctate. Meso- and metatarsi slightly shorter than in male; tarsomeres slender and elongate. Dimensions. Male/female. Total length (including mandibles) = /50.0 mm; prothoracic length = /7.0 mm; prothoracic width between apices of anterior angles = /10.4 mm; prothoracic width between apices of posterior angles = /12.0 mm; humeral width = /17.3 mm; elytral length = / 28.8 mm.

29 228 THE COLEOPTERISTS BULLETIN 67(3), 2013 Figs Prionacalus iphis. 76) Male, dorsal view; 77) Male, ventral view; 78) Male, head, dorso-lateral view; 79) Male, mandibles; 80) Male, protarsus; 81) Male, mesotarsus; 82) Male, metatarsus; 83) Male, protibia; 84) Female, dorsal view; 85) Female, ventral view; 86) Female, mandibles; 87) Female, metatarsus.

30 THE COLEOPTERISTS BULLETIN 67(3), Geographical Distribution. Ecuador (Waterhouse 1872) and Peru (new country record). Material Examined. ECUADOR: Male, ex Deyrolle Collection (no other data) (ZMHB); male, ex Deyrolle Collection (no other data) (IRSN); male, ex Candèze Collection (no other data) (IRSN); route between Loja and Cuenca, km 20 (2,500 m), male, II , [no collector indicated] (ZKCO). Manabí: Portoviejo, male, ex Nonfried Collection (no other data) (IRSN). Loja: Cordillera del Cóndor (peak above Loja; 3000 m), male, 19.IX.1905, F. Ohaus col. (ZMHB). PERU: Male, ex Moffarts Collection (no other data) (IRSN). Bolivar: 1 male, 1, female, III.2007, [no collector indicated] (ZKCO). Cañar: 11 km N Zhud (2850, on the Road to Canton Guamote), 1 male, 1 female, 28.I.1987, N. Krable col. (ZMUC). Types and Type Localities. Of Prionacalus iphis (Figs ): See Types and Type Locality for P. cacicus. The website of the BMNH (2013) records only one type of Prinocalus [sic] iphis deposited there (sex not indicated): holotype. As mentioned above, it is not possible to know if the third specimen mentioned by White (1845) corresponds to P. cacicus or P. iphis. Thus, the holotype of P. iphisdeposited in the BMNH is considered here the lectotype of the species [designation by Waterhouse (1872)]. Of Prionocalus [sic] buckleyi (Figs ): Described based on two males and a single female, all from Ecuador ( Yerba buena ). All syntypes should be deposited in the BMNH, because Waterhouse (1872) indicated this ( Brit. Mus. ). There are currently two syntypes deposited at BMNH. According to the Directory of Cities and Towns in World ( ), there are two places with names equal or similar to Yerba buena in Ecuador. These places are in different provinces: Yerbas Buenas in Cañar, and Yerbabuena in Azuay. Therefore, the exact type locality is not known. The website of the BMNH (2013) records two syntypes deposited in the institution (unknown sexes). However, there is no doubt that Waterhouse had, at least, two males. He described a male with 28 lines length (about mm) but also another male with 18 lines length (about mm). We designate as the lectotype of P. buckleyi the specimen (Figs. 112, 113) with the following labels (Fig. 114): 1. White [Handwritten]: yerba buena 2. White with greenish circle [Printed]: Syntype 3. White [Handwritten]: Prionocalus [sic] Buckleyi, /. C.Waterh./(Type.) 4. Red and yellow [Printed; added by us]: LECTOTYPE / Prionacalus buckleyi Of Prionacalus emmae: Described based on a single male from Ecuador (between Las Palmas and Chapacoto), deposited in the ZMHB. Chapacoto is in the Province of Bolívar. According to Directory of Cities and Towns in World ( ), there are six places named Las Palmas in five Ecuadorian provinces, none of them in Bolívar. Kolbe (1902) recorded (translation): was on the crest of the Western Cordillera of Ecuador Auguste (1902) referred to the locality in connection with an undescribed species of Palicourea Aubl., 1775 (Rubiaceae) (translation): Between Las Palmas and the top of the pass toward Chapacoto, on the way to Guaranda; forested slope of the Western Cordillera (Ecuador). Thus, it is possible to affirm that the province where the specimen was collected is Bolívar. Of Prionocalus [sic] whitei (Figs ): Described based on a single male from Ecuador (Porvenir), deposited in the BMNH. According to Directory of Cities and Towns in World ( ), there are many places named Porvenir in Ecuador. These places are in different provinces: Francisco de Orellana, Manabí, Pastaza, Guayas, Esmeraldas, and Pichincha. Thus, it is not possible to know exactly where the type locality of P. whitei is. The holotype has two labels added after Waterhouse (1900): Type and Syntype. Without a doubt, the addition of the Syntype. label was in error, because Waterhouse (1900) gave a single measure: Long. 50 mm. Remarks. Waterhouse (1880) stated for Prionacalus buckleyi (the name of the genus was correctly spelled by the author, and not as Prionocalus as was misspelled in the original description of this species): In the Prionidae, the specimens of Prionacalus Buckleyi, W.,taken by Mr. Buckley differ immensely in size and development, in the same way as is seen in P. cacicus and P. a t y s ; and the sculpture of the thorax varies also somewhat in all the species. I have seen one small example of P. Buckleyi. This reference is not mentioned in Monné s (1995, 2006) catalogues. Lameere (1910) synonymized four species with P. buckleyi (translation): I have not seen the type of P. Whymperi Bates; the species is constituted of a male slightly differing of P. Buckleyi, according to Bates himself, but also on a female in which the elytra substantially exceed the apex of abdomen. I have before me one male of P. Buckleyi which also offers this feature, but I cannot see more than an individual variation. ; P. W h i t e i C. O. Waterh. also was based on a single male closely resembling the type of P. Gunteri, also having the elytra separately rounded at apex, but the abdomen is quite rough and the sculpture of elytra is uniform; the tubercle behind eyes is

31 230 THE COLEOPTERISTS BULLETIN 67(3), 2013 exceptionally thick and obtuse. ; P. E m m a e Kolbe was described on a single male that seems to be just a P. Buckleyi with reddish appendices. Afterwards, Lameere (1913, 1919), without explanation, recorded P. whiteias a variety of P. buckleyi, but kept P. whymperi and P. emmae as its synonyms. Thus, Lameere (1913) revalidated the former species (not in 1910 as pointed out by Monné and Giesbert 1994 and, Monné 1995, Monné 2006). According to the ICZN (1999: ), Lameere (1913) is not the author of the species (P. whitei), but since he considered it a variety of P. buckleyi, it should be considered a subspecies. It is important to note that Quentin and Villiers (1983) and Hüdepohl (1985) mentioned only P. buckleyi, without a subspecies. Quentin and Villiers (1983) revalidated P. whymperi. Hüdepohl (1985), in his keys to the species, did not mention P. w h y m p e r i (probably following the synonymy by Lameere (1910)), and suggested a synonymy between P. trigonodes and P. woytkowskii (he did not formalize the synonymy). We agree with the synonymy of P. emmae with P. buckleyi (and thus, with P. i p h i s ). Although Kolbe (1902) had pointed out that P. emmae differs in many aspects from P. buckleyi, all characters mentioned by him are variable in all species of Prionacalus (e.g., mandibular shape, color, punctation). Based on the original description and drawing, we can affirm that Lameere (1910) was correct in proposing the synonymy. Since Lameere (1910), P. emmae has been mentioned as having been published in However, although Sechsundvierzigster Band (1901) [forty-sixth volume (1901)] is recorded on the book cover of the magazine, it is also recorded as below: Viertes Heft: (IV), / Mit 3 Tafeln und 4 Textfiguren / Ausgegeben Mitte Februar 1902 [Fourth Issue: (IV), / With 3 plates and 4 text figures/ Issued mid February 1902]. Thus, the record of Kolbe, 1901 is an error that needs to be corrected (see, for example, in Monné 2006). We believe that Lameere (1913) considered Prionacalus whitei a variety of P. buckleyi because the holotype has a different elytral apex (more or less jointly rounded in P. buckleyi, and individually rounded in P. whitei). However, the elytral apex is variable in species of Prionacalus. The same variation occurs with the holotype of P.emmae, also considered by Lameere (1913) as a variety of P.buckleyi. We think that P. w h y m p e r i is distinct from P. buckleyi (and so, from P. i p h i s ) and therefore agree with the revalidation by Quentin and Villiers (1983). Lameere (1910) pointed out the following variations in P. buckleyi (translation): 1 on the coloring of its appendages that are distinctly dark or reddish; 2 on the development more or less pronounced of its cephalic carinae and tubercle behind eyes; 3 on the extending more or less large of the roughness of pronotum that sometimes has smooth spaces; 4 on the sculpture of elytra a little more or a little less rough; 5 on the projection of elytral humeri; 6 on the shape of sutural angle, the elytra are sometimes together rounded at apex, sometimes individually rounded; 7 on the length and shape of elytra that surpass more or less the abdomen; 8 on the punctation on abdomen which is more or less rough. Lameere (1910) used the expression more or less in the sense of slighter or stronger and not as about. We infer this because he also stated: I have not seen the type of P. Whymperi Bates; the species is constituted on a male slightly different from P. Buckleyi according to Bates himself, but also on a female with elytra notably surpassing the apex of abdomen. All those variations occur in P. buckleyi (= P. iphis), and also they were observed by us in nearly all other species of the genus. Lameere (1910) wrote on P. iphis(translation): the punctation of abdomen is fine. However, the punctation on the abdomen of P. iphisis somewhat variable; this was also recorded by Lameere (1910) when he wrote on P. buckleyi(translation): from abundant to moderately sparse; from moderately fine to distinctly coarse. Thus, this difference does not allow for separation of the species. The same is true with other characters pointed out by Lameere (1910) to define P. i p h i s (translation): the antennae slightly surpass elytral apex; the humerus is slightly dentate and little projected In his key to species, Lameere (1910) recorded that in P. iphis (translation): Elytra dentate at suture. He also noted that in P. buckleyi (translation): This species differs from P. iphis by its elytra not angled at the suture; its humerus is very spinose and very projected; its antennae surpass the apex of the body in male, and reach the posterior third of elytra in female. All these differences are quite variable in P. iphis. However, the most important character here is the shape of the elytral apex (angled). Lameere affirmed that this is a character seen in P. buckleyi.thischaracter was later used by other authors to separate P. iphisin their keys (translations): Elytra dentate at suture (Quentin and Villiers 1983); Tips of elytra dentate at suture (Hüdepohl 1985). Although the elytral apex of P. iphis can be dentate (a variation that also occurs in all other known species of the genus), the holotype does not have the elytral apex dentate, it is angled. Apparently, Quentin and Villiers (1983) and Hüdepohl (1985) incorrectly interpreted Lameere (1910). Lameere stated that the elytral apex is angled at the suture, not dentate.

32 THE COLEOPTERISTS BULLETIN 67(3), We examined specimens of P. i p h i s in which the humeral spines were almost absent and with ventrites coarsely and abundantly punctate, and others with a very distinct and long humeral spine and ventrites distinctly finer punctate. These variations found in the specimens lead us to conclude that P. buckleyi is only a variation of P. iphis. Furthermore, the length of the antennae is affected by the length of the elytra. Comparing the lectotype of P. iphis and a syntype male of P. buckleyi, itis evident that the former has longer elytra and, therefore, its antennae are proportionally shorter. Prionacalus uniformis Waterhouse, 1900 (Figs ) Prionocalus uniformis Waterhouse 1900: 505, 1 fig.; Quentin and Villiers 1983: 225 (key); Hüdepohl 1985: 121 (key). Psalidognathus (Prionocalus) uniformis; Lameere 1910: 375, 1913: 65 (catalogue), 1919: 121 (checklist); Blackwelder 1946: 555 (checklist). Prionacalus uniformis; Monné and Giesbert 1994: 15 (checklist); Monné 1995: 61 (catalogue), 2006: 84 (catalogue); Komiya 2001: 30 (fig. 4), 31, 32 (key); Di Iorio 2003: 3 (distribution); Monné and Hovore 2005: 19 (checklist), 2006: 18 (checklist); Jeniš 2010: 19 (syntype male), 111 (male); Monné et al. 2012: 7 (checklist). Redescription. Lectotype male (Fig. 124). Integument blackish with some parts dark brown; scape, pedicel and basal 1/2 of antennomere III dark brown; antennomeres IV-XI reddish, slightly lighter on distal antennomeres; palpi reddish; elytra blackish with areas slightly reddish, mainly around basal sutural 1/2; femora dark brown with blackish areas; tibiae blackish on base, gradually becoming lighter towards reddish apex; tarsi reddish, with part of claws blackish. Head: Coarsely punctate-vermiculate between cephalic carinae; area between apex of cephalic carinae and pronotum coarsely, abundantly, confluently punctate; area between cephalic carinae with well-marked, longitudinal furrow from clypeus to after posterior ocular edge; distal margin of vertex (close to abrupt declivity) centrally distinctly emarginate. Antennal tubercle coarsely, sparsely punctate on base, gradually becoming smoother towards apex; distinctly far from each other (distance between them equal to almost twice width). Cephalic carinae distinct, moderately elevated from base to apex; convergent from antennal tubercle to slightly after eyes; apex not elevated, not forming triangular tubercle. Area behind eyes with abundant small asperities. Lateral tubercle of head large, rounded at apex. Distal area of clypeus distinctly concave. Distance between upper ocular lobes equal to about 3.0 times longest width of 1 lobe. Gena rounded at apex, without downward projection. Submentum coarsely, abundantly, confluently punctate. Mandibles shorter than head; surface coarsely, abundantly punctate, gradually finer towards apex. Last maxillary and labial palpomeres notably enlarged towards apex. Antennae reaching distal 1/3 of elytra; scape distinctly enlarged from base to apex, coarsely, confluently punctate on basal 1/2, gradually finer, sparser towards apex; antennomeres III-IV rounded at apex on both sides; antennomeres V-X dentate at outer distal angle. Thorax: Latero-anterior angles of prothorax projected forward, rounded; latero-posterior angles of prothorax distinctly rounded, not projected; lateral margin with 3 jointly protracted teeth, the anterior tooth smaller than others. Pronotum centrally with distinct, transverse depression; surface coarsely, confluently punctate; disc with 1 small, elliptical, impunctate, shiny callosity on each side close to anterior margin. Elytra coarsely vermiculate throughout; almost 3.0 times longer than prothorax; apex individually rounded. Apical tooth of humeral projection small. Abdomen: Not surpassing elytral apex. Ventrites (Fig. 125) with coarse punctures, moderate on I and becoming more abundant towards V. Legs: Apex of metafemora distinctly surpassing elytral apex. Ventral sulcus of protibiae distinct from near base to distal constriction. Protarsomeres I and II short, moderately wide; protarsomeres I-III with very small, apical denticle; metatarsi slender. Dimensions. Male. Length mm (Waterhouse 1900). Geographical Distribution. Peru. Material Examined. PERU: San Pablo, lectotype male, P. O. Simons col. (BMNH). Types and Type Locality. This species was apparently described based on two syntype males from Peru. According to the website of the BMNH (2013), there is only one syntype deposited there. EHN examined this specimen and confirmed that it is the only specimen in that collection. We designate as the lectotype of P. u n i f o r m i s the specimen (Figs. 124, 125) with the following labels (Figs. 126, 127): 1. White with red circle [Printed]: Type 2. White with red circle [Printed]: Type 3. White with greenish circle [Printed]: Syntype 4. White [Handwritten]: San Pablo / Peru metres / P. O. Simons by. 5. White [Handwritten]: Prionocalus uniformis (Type) Waterh. 6. White [Handwritten]: upper timber line 7. Red and yellow [Printed; added by us]: LECTOTYPE / Prionacalus unifromis

33 232 THE COLEOPTERISTS BULLETIN 67(3), 2013 Figs Prionacalus species. 88) P. demelti, holotype male, dorsal view; 89) P. demelti, holotype, labels; 90) P. demelti, allotype female, dorsal view; 91) P. d em el ti, allotype, labels; 92) P. trigonodes, holotype male, frontal view; 93) P. trigonodes, holotype, dorsal view; 94) P. trigonodes, holotype, ventral view; 95) P. trigonodes, holotype, labels.

34 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus whymperi. 96) Lectotype male, dorsal view; 97) Lectotype, ventral view; 98) Lectotype, frontal view; 99) Lectotype, labels; 100) Paralectotype female, dorsal view; 101) Paralectotype, ventral view; 102) Paralectotype, labels.

35 234 THE COLEOPTERISTS BULLETIN 67(3), 2013 Figs Prionacalus species. 103) P. atys, holotype male, dorsal view; 104) P. atys, holotype, ventral view; 105) P. a t ys, holotype, labels; 106) P. cacicus, syntype male, dorsal view; 107) P. cacicus, syntype,ventral view; 108) P. cacicus, syntype, labels.

36 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus species. 109) P. gunteri, holotype male, dorsal view; 110) P. gunteri, holotype, ventral view; 111) P. gunteri, holotype, labels; 112) P. buckleyi, lectotype male, dorsal view; 113) P. buckleyi, lectotype, ventral view; 114) P. buckleyi, lectotype, labels.

37 236 THE COLEOPTERISTS BULLETIN 67(3), 2013 Figs Prionacalus species. 115) P. iphis, lectotype male, dorsal view; 116) P. iphis, lectotype, ventral view; 117) P. iphis, lectotype, labels; 118) P. whitei, holotype male, dorsal view; 119) P. whitei, holotype, ventral view; 120) P. whitei, holotype, labels.

38 THE COLEOPTERISTS BULLETIN 67(3), Figs Prionacalus species. 121) P. simonsi, syntype male, dorsal view; 122) P. simonsi, syntype male, ventral view; 123) P. simonsi, syntype male, labels; 124) P. uniformis, lectotype male, dorsal view; 125) P. uniformis, lectotype male, ventral view; ) P. uniformis, lectotype male same, labels.

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