Beneath the bark - Complicated taxonomy: Revision of the darkling beetle genus Phrenapates Gray, 1832 (Tenebrionidae: Phrenapatinae)
Abstract
Rincon, Angela, Lumen, Ryan, Kamiński, Marcin J., Smith, Aaron D. (2025): Beneath the bark - Complicated taxonomy: Revision of the darkling beetle genus Phrenapates Gray, 1832 (Tenebrionidae: Phrenapatinae). European Journal of Taxonomy 1012: 49-80, DOI: 10.5852/ejt.2025.1012.3025, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3025/13587
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49 European Journal of Taxonomy 1012: 49–80 https://doi.org/10.5852/ejt.2025.1012.3025 europeanjournaloftaxonomy.eu ISSN 2118-9773 2025 · Rincon A. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0) Received: 28 February 2025 • Accepted: 30 June 2025 • Published: 29 August 2025 Topic editor: Tony Robillard • Section editor: Maxwell Barclay • Desk editor: Eva-Maria Levermann Research article urn:lsid:zoobank.org:pub:62D69A2D-D84E-4D80-B563-981969B6EB7F Beneath the bark – Complicated taxonomy: Revision of the darkling beetle genus Phrenapates Gray, 1832 (Tenebrionidae: Phrenapatinae) Angela RINCON 1,* , Ryan LUMEN 2 , MarcinJ.KAMIŃSKI3 & Aaron D. SMITH 4 1,4 Department of Entomology, Purdue University, West Lafayette, USA. 2,3 Zoological Museum, Museum and Institute of Zoology, Polish Academy of Sciences, Warszawa, Poland. * Corresponding author: [email protected] 2 Email: [email protected] 3 Email: [email protected] 4 Email: [email protected] Abstract. A revision of the Neotropical darkling beetle genus Phrenapates Gray, 1832 (Tenebrionidae: Phrenapatinae) is conducted. The genus previously contained six species ranging from Central to South America. Nearly 350 specimens of Phrenapateswereexaminedtodefinemorphologyandevaluate previouslypublishedspecies.Thegenusisredescribedwithitstaxonomichistoryandclassification summarized. Two new species are described: Phrenapates gilloglyi Rincon & Smith sp. nov. from the Cerro Jefe Mountain located within Cerro Azul, Panama; and Phrenapates fortunaensis Rincon & Smith sp. nov. from the Fortuna Forest Reserve in Panama. A replacement name, Phrenapates erratus Rincon, Lumen&Kamińskinom.nov.,isintroducedforPhrenapates latreillei Gebien, 1910 to avoid homonymy with Phrenapates latreillei Lacordaire, 1859. Lectotypes are designated for Phrenapates bennettii Gray, 1832, P. dux Gebien, 1910, P. educator Gebien, 1910, P. erratus, P. latreillei, and P. ohausi Gebien, 1910, to stabilize species concepts. Phrenapates mandibularis Gebien 1910 is treated as a junior synonym of Phrenapates latreillei Lacordaire, 1859. As a result, a total of eight species is now recognized (P. bennettii, P. ohausi, P. dux, P. latreillei, P. educator, P. erratus, P. gilloglyi, and P. fortunaensis). Distribution maps and a key to the species of Phrenapatesareprovided.Dissectionsconfirmedthat unlike most other Tenebrionidae, female Phrenapates lack well-sclerotized ovipositors, with only coxite 4 and paraproct remaining for oviposition. This reduction is potentially attributed to their unusual life history, as Phrenapates live in subsocial colonies inside dead wood and may not need to inject eggs into substrate. Keywords. Darkling beetles, Neotropical, Panama, new species, nomen novum. RinconA.,LumenR.,KamińskiM.J.&SmithA.D.2025.Beneaththebark–Complicatedtaxonomy:Revisionof the darkling beetle genus Phrenapates Gray, 1832 (Tenebrionidae: Phrenapatinae). European Journal of Taxonomy 1012: 49–80. https://doi.org/10.5852/ejt.2025.1012.3025
European Journal of Taxonomy 1012: 49–80 (2025) 50 Introduction Phrenapatinae Solier, 1834 is a diverse subfamily of Tenebrionidae Latreille, 1802 containing 25 genera and ca 150 described species of saproxylic darkling beetles distributed across the Afrotropical, Australasian, Indomalayan, Nearctic, Neotropical, and Palearctic regions (Watt 1974; Matthews et al. 2010; Rosas-Ramos 2020; Bouchard et al. 2021). To accompany their subcortical lifestyle, phrenapatines express an array of cuticular head armaments and variable mandible morphology. Current taxonomy has beenfocusedonalphalevelcontributions(Schawaller&Bouchard2019;Iwan&Raś2020;Jianget al. 2020; Iwan et al. 2023). The type genus for the subfamily, Phrenapates Gray, 1832, was last treated by Gebien (1911), who also provided additional descriptions and illustrations. The distribution of Phrenapates spans from Central to South America (Doyen 1979). They largely live in colonies inside logs and stems of dead trees (ex. Bombax Linnaeus, 1753 also known as silk-cotton trees), with larvae developing within lateral chambers and depending upon the frass generated by adults (Ohaus 1909; Watt 1974). Phrenapates superficiallyresemblebeetlesinthefamilyPassalidaeLeach, 1815 (Fig. 1) who also exhibit parental feeding behavior and live in subcortical colonies (Shuster & Shuster 1985, 1997). The bauplan of Phrenapates resembles that of lagriioid Tenebrionidae (see Tschinkel&Doyen1980),withthedistinguishingcharacterbeingthefirstsegmentoftheantennaebeingas long as the following 3, as well as the last 3 segments forming a club and having elongated mouthparts (e.g., projecting mandibles, galea longer than lacinia) (Solier 1834; Watt 1974; Doyen & Lawrence 1979). GeorgeGray(inGriffith&Pidgeon1832)providedthefirstvalidspeciesdescriptionofthegenus using an unpublished account by William Kirby (Phrenapates bennettii Gray, 1832). In his subsequently published work, Kirby (1837) described the similarity of Phrenapates to Passalidae and Bostrichidae Latreille, 1802, using the following characters: body plan similar to Passalidae, spurious joint between the claws of the tarsi, and all tibiae possessing spines. Solier (1834) placed Phrenapates in a new tribe Phrépatides within the Heteromeres (now Tenebrionidae), using the number of antennomeres (11) and the number of tarsomeres on the fore, mid, and hind tarsus (5-5-4) as uniting characters. In 1859, Lacordaire published a catalog of Tenebrionidae mentioning Kirby’s description of P. bennettii and providing a description for P. latreillei Lacordaire, 1859, erroneously attributing it to Dejean. Gebien in 1910 reFig. 1. Convergent morphology of Passalidae Leach, 1815, and Phrenapates Gray, 1832. A. Passalidae Leach, 1815 (MIZPAN). B. Phrenapates bennettii Gray, 1832 (MIZPAN).
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 51 described P. bennettii andfiveadditionalspecies,bringingthetotalspeciescounttosix.In1911,Gebien provided further descriptions and illustrations in a second publication. Watt in 1974 elevated Phrenapatinae to subfamily status and placed Phrenapates into the tribe Phrenapatini Solier, 1834. To reassess, organize, and identify current and undescribed species, 346 specimens were loaned from eleven institutions, including type material from all previously known species. A morphological revision was conducted to test the taxonomic stability of the genus, as well as to describe new species. The last treatment of Phrenapates was over 100 years ago, and since its original description(s), more material has become available to evaluate species concepts, making a revision of the type genus of the subfamily Phrenaptinae (and tribe Phrenapatini) possible. Material and methods Institutional abbreviations The following collections graciously loaned specimens for this study. AAAG = Alan and Anita Gillogly (private coll.), Caldwell, Idaho; Alan Gillogly AMNH = American Museum of Natural History, New York, USA; Lee Herman BMNH = The Natural History Museum [formerly British Museum (Natural History)], London, UK; Max Barclay CASC = California Academy of Sciences, San Francisco, California, USA; Christopher Grinter FMNH = Field Museum of Natural History, Chicago, Illinois, USA; Julia Snyder MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA; Crystal Maier MIZPAN = Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw, Poland; Dariusz Iwan MNHN = Muséum national d’Histoire naturelle, Paris, France: Antoine Mantilleri NHMB = Naturhistorisches Museum, Basel, Switzerland; Christoph Germann PERC = Purdue Entomological Research Collection, Purdue University, West Lafayette, Indiana, USA; Aaron D. Smith SEMC = Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA; Kirsten Jensen TAMU = Texas A & M University, College Station, Texas, USA; Karen Wright This revision is based on morphological and georeference data acquired from 346 adult specimens of Phrenapates. Of these, subsets of each species were measured for size (total of 105 specimens). DissectionfollowedthemethodologyofIwan&Kamiński(2016).Specimensweresoakedinawarm KOH solution to soften connective tissue before dissection. Morphological terminology follows that of Gebien (1911), Doyen (1966), and Matthews et. al(2010).Materialidentifiedviacomparisonwith type material had their mouthparts removed and mounted on cards or points. The KOH method was also used to dissect reproductive structures. Dissected tissue was then included in a vial or paper point/ card on the pin beneath the specimen. Images of specimens were taken using a Passport Imaging system (https://www.visionarydigital.com), and mounted images assembled using Helicon Focus ver. 5.3 (https://www.heliconsoft.com). Backgrounds were cleaned in Adobe Photoshop CS6. Measurements were taken digitally using the ruler tool in Photoshop on images with known measurements based onthecamerabody,lens,andmagnificationused.Smallerpartsofspecimenswerephotographed, montaged, and measured using a Leica M165 stereo microscope with the accompanying Leica software (https://www.leica-microsystems.com). Body length was measured along the midline from the apex of mandibles to the apex of elytra. Body width was measured across the widest point of the elytra. Color wasdeterminedunderfiberopticilluminationandfromimages.Distributionaldatafromspecimenlabels
European Journal of Taxonomy 1012: 49–80 (2025) 52 were used to generate maps using QGIS ver. 3.16, with vector layers downloaded from the Natural Earth web page (https://www.naturalearthdata.com). Results Themajorityofspecimensexaminedweremisidentifiedorlackedspecieslevelidentifications.Upon examination of pinned material and comparison of type and more recently collected specimens, it was clear that most differences between species are restricted to the head. Further, the original diagnoses and descriptions were often vague or inaccurate (e.g., uninformative reproductive morphology at the species level or subtle differences between similar taxa such as Phrenapates educator vs P. erratus Rincon, Lumen&Kamińskinom.nov.).Assuch,anewgenericdefinitionoutliningthevariationofmorphology within the group is provided in the taxonomy section below, as well as newer treatments at the species levelforpreviouslydescribedspecies.Specifically,typedesignationsforpreviouslydescribedspeciesare provided where necessary (e.g., lectotypes), along with brief diagnostic redescriptions as well as focused diagnoses to provide stability to the revised species concepts presented here. Taxonomy Class Insecta Linnaeus, 1758 Order Coleoptera Linnaeus, 1758 Suborder Polyphaga Emery, 1886 Family Tenebrionidae Latreille, 1802 Subfamily Phrenapatinae Solier, 1834 Tribe Phrenapatini Solier, 1834 Genus Phrenapates Gray, 1832 Phrenapates Gray, 1832: 91. Phrepates Solier, 1834: 488 [incorrect subsequent spelling of genus name]. Type species Phrenapates bennettii Gray, 1832; by monotypy. Diagnosis In terms of body size, Phrenapates is the largest genus in the subfamily by a large margin (18–39 mm long vs Archaeoglenini Watt, 1974 < 5 mm and Penetini Lacordaire, 1859 < 10 mm). Additionally, Phrenapates is the only genus in the subfamily to possess both clypeal horns and supraorbital tubercles in males and females; other examples of armature in the subfamily include projections on the head (e.g., Peneta Lacordaire, 1859) and / or mandibles (Molion Champion, 1886). The only other genus in the tribe Phrenapatini, Delognatha Lacordaire, 1859, can be differentiated based on the following characteristics: clypeal horns and supraorbital tubercles absent, smaller (less than 8 mm), and mandibles less projected (< the length of the head). Archaeoglenines can be differentiated from Phrenapates by the following characters: extremely small size (< 5 mm), procoxal cavity closed laterally by mesoand metaventrites, mesotrochantin not visible; prothorax with distinct anterolateral antennal cavities (Iwan et al. 2015). Phrenapates can be differentiated from penetines as they have mesocoxae with exposed trochantins and prominent mandibles (Watt 1974; Doyen & Lawrence 1979). Etymology No etymology was given in the original description. It potentially derives from the Greek roots ‘Phren-’ (‘place of thought’ / ‘head/mind’) and ‘-apata’ (‘deceitful’ / ‘cheat’), possibly referencing the head’s morphological similarity to passalids.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 53 Redescription MeasureMents. Length 18–39 mm, width 6–13 mm, specimens measured (n = 105). Color nitidus dark brown to black. Body sub-cylindrical. H ead . Antennae clavate 11-segmented with last three antennomeres forming club, each with sensilla on apices. Subgenal ridge slightly dilated distally, tapering near mentum. Mandibles protruding and tridentate, equal to or greater than length of head, medial edge with or without teeth, inner margin crenulate or not. Mandibles with or without small setae on dorsal surface. Galea long and cylindrical with cluster of bristles situated near midpoint or toward apex. Mentum rectangular, slightly widening towards apex, strongly to weakly sculptured, apex with medial notch or not; lateral apices produced or not (Fig. 2). Clypeus apex dilated or not, all species with clypeal horn in both sexes. Clypeal horn variously shaped (Fig. 3) and occasionally slightly twisted or bent to left or right. Supraorbital tubercles present or absent. Eyes generally larger in males than females, roughly circular, undivided, and slightly protuberant. ProtHorax. Pronotum rectangular, micro-punctate, lateral margin carinate. Posterior of head capsule tucked slightly beneath anterior margin. PterotHorax. Scutellum sub-triangular. Elytral striae with large punctures evenly spaced and interstices with small irregularly spaced punctures. Macropterous. Metaventral groove shallow and broad. Legs. Tibiae armed with 0–5 spines, apex with four pointed spines where tarsi attach. Sparse setae on apices of tibiae and on tarsi. Protibiae with cluster of long apical setae. Pulvilla with four setae. abdoMen. Glabrous, shining, and impunctate. Hind margins of abdominal ventrites three and four with membranes exposed. Aedeagus bipartite, parameres slightly dilated apically in lateral view, rounded apically in dorsal view with 4–5 setae on each paramere (Fig. 4). Median lobe exits ventrally. Ovipositor reduced to sclerotised coxite 4 and weakly sclerotised paraproct (Fig. 5). Remarks For information on larvae and pupae, see descriptions by Doyen & Lawrence (1979) and pupal descriptions by Bouchard (2019). Status MemberofthesubfamilyPhrenapatinaeintribePhrenapatini,whichisunifiedbythemesocoxaelacking exposed trochantins and the possession of prominent mandibles (Watt 1974). Phrenapatini includes two genera, Phrenapates and Delognatha. Species included Phrenapates bennettii Gray, 1832, P. dux Gebien, 1910, P. educator Gebien, 1910, P. erratus Rincon, Lumen&Kamińskinom.nov., P. fortunaensis Rincon & Smith sp. nov., P. gilloglyi Rincon & Smith sp. nov., P. latreillei Lacordaire 1859, P. ohausi Gebien, 1910. Distribution (Figs 6–7) Central America: Costa Rica, Guatemala, Honduras, Nicaragua, Panama; South America: Bolivia, Brazil, Chile, Colombia, Ecuador, Guyana, Peru, Venezuela.
European Journal of Taxonomy 1012: 49–80 (2025) 54 Fig. 2. Different menta morphologies of the species of Phrenapates Gray, 1832. A. Phrenapates bennettii Gray, 1832, lectotype, ♂ (Teneb_base 23170, BMNH). B. Phrenapates gilloglyi Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23367, PERC). C. Phrenapates dux Gebien, 1910, (Teneb_base 23242, PERC). D. Phrenapates erratus Rincon, Lumen & Kaminski nom. nov.,lectotype,♂(Teneb_base23168, NHMB). E. Phrenapates mandibularis Gebien 1910 [Phrenapates latreillei Lacordaire],holotype,♂ (Teneb_base 23167, NHMB). F. Phrenapates educator Gebien,1910,lectotype,♂(Teneb_base23175, NHMB). G. Phrenapates fortunaensis Rincon & Smith sp. nov.,holotype,♂(Teneb_base23482,PERC). H. Phrenapates ohausiGebien,1910,lectotype,♂(Teneb_base22810,BMNH).
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 55 Fig. 3. Clypeal horn morphology, lateral views. A. Phrenapates ohausi Gebien, 1910 (Teneb_base 23212, CASC). B. Phrenapates bennettii Gray, 1832 (Teneb_base 22972, AAAG). C. Phrenapates erratus Rincon,Lumen&Kamińskinom.nov.,lectotype,♂ (Teneb_base 23168, NHMB). D. Phrenapates dux Gebien, 1910 (Teneb_base 23215, AAAG). E. Phrenapates fortunaensis Rincon & Smith sp. nov., holotype,♂(Teneb_base23482,PERC).F. Phrenapates gilloglyiRincon&Smithsp.nov.,holotype,♂ (Teneb_base 23367, PERC). G. Phrenapates latreillei Lacordaire, 1859, ♂(Teneb_base22740,AAAG). H. Phrenapates educator Gebien, 1910 (Teneb_base 23286, MNHM)
European Journal of Taxonomy 1012: 49–80 (2025) 56 Fig. 4. Phrenapates Gray, 1832, adeagi, dorsal and lateral views. A–B. Phrenapates bennettii Gray, 1832,♂(Teneb_base23186,NHMB).C–D. Phrenapates dux Gebien, 1910, lectotype, ♂ (Teneb_base 23166, NHMB). E–F. Phrenapates educator Gebien, 1910, lectotype, ♂ (Teneb_base 23175, NMHB). G–H. Phrenapates mandibularis Gebien 1910 [Phrenapates latreillei],holotype,♂(Teneb_base23167, NHMB). I–J. Phrenapates ohausi Gebien,1910,♂(Teneb_base23165,NHMB).
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 57 Fig. 5. Female terminalia of two species of Phrenapates Gray, 1832. A. Phrenapates bennettii Gray, 1832 (MIZPAN). B. Phrenapates dux Gebien, 1832 (MIZPAN). Abbreviations: an = anus; bp = baculus of paraproct; c4 = coxite 4; gt = genital tubes; p = paraproct. Fig. 6. Distribution of the genus Phrenapates Gray, 1832, Phrenapates bennettii Gray, 1832, Phrenapates educator Gebien, 1910, and Phrenapates ohausi Gebien, 1910.
European Journal of Taxonomy 1012: 49–80 (2025) 64 P. duxisespeciallyuniquewithinthegenus,beingconstrictedatthebase,dorsallyflattenedonthetop, and having an angular arc. Other species without supraorbital tubercles are differentiated by clypeal horns with strong, circular arcs (P. latreillei), or with weak arcs (P. educator). The lateral apices of the mentum are produced in P. dux, rather than rounded of P. ohausi. The dorsal ridge next to the clypeus is not dilated as in P. educator. Mandibles without the crenulations seen in P. bennettii. Type material Lectotype (designated here) ECUADOR•♂;(greenlabel)“O.Ecuador./Macas//Phrenapates / dux / X Geb (red label) Cotype! / No. 166 // Sammlung / H Gebien // (blue label) Tenebrionid Base / Aaron D. Smith / Catalog # 23166”; Teneb_base 23166; NHMB. Fig. 12. Phrenapates dux Gebien, 1910, lectotype, ♂ (Teneb_base 23166, NHMB), dorsal (top) and lateral (bottom) views.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 65 Other material examined BOLIVIA • 1 spec.; Yungas de La Paz; Teneb_base 23191; NHMB • 1 spec.; Yungas de LePaz, 1000m; H. Rolle Berlin S.W.N. leg.; Teneb_base 23184; NHMB. CHILE • 1 spec.; [northern] Chili; 29 Dec. 1973; [C]. G. Minet leg.; Teneb_base 23259; NHMB. ECUADOR • 1 spec.; Napo, 3 km E of Huaticocha; 0°45.22' S; 77°27.84 W; 19 Aug. 1997; A.R. Gillogly leg.; Teneb_base 22949; AAAG • 2 specs; Napo Prov. Archidona, Rio Hollin; 21 Dec. 2001; P. Bravo and E. Vasquez leg; collection R. Diaz; Teneb_base 23214 to 23215; AAAG • 3 specs; Zatzayacu Oriente; 10 Nov. 1932; Teneb_base 23222 to 23224; AMNH • 5 specs; El Partidero; 17–27 Nov. 1935; Teneb_ base 23225 to 23228, 23233; AMNH • 1 spec; Zatzayacu Oriente; 10 Nov. 1932; Teneb_base 23229; AMNH • 1 spec; [East] Ecuador Jivaria; 17 Nov. 1905; Ohaus leg.; Teneb_base 22809; BMNH • 2 specs; Quito F4724; Teneb_base 23220 to 23221; CASC • 1 spec.; Macas er.; Teneb_base 23275; CASC • 1 spec.; Guayaquil SA; Teneb_base 23199; CASC • 4 specs; 3–12 m N of Puyo, Napo Pastaza; 935 m a.s.l.; 9 Nov. 1955; E.I. Schlinger and E.S. Ross leg.; Teneb_base 23192 to 23195; CASC • 1 spec.; Lumbaqui; 23 Sept. 1977; Teneb_base 23201; FMNH • 3 specs; Arajuno; Oct. 1960; L.E. Pena coll. acc. 17-422; Teneb_base 23196 to 23198; FMNH • 1 spec.; Pastaza Jibaria Shurupe; 7 Nov. 1987; Mike Huybensz leg.; Teneb_base 23202; FMNH • 1 spec.; 1853; Museum Paris Chimborazo, and Pichincha MANDEVILLE; Teneb_base 23273; MNHN • 1 spec.; 1922; Teneb_base 23274; MNHN • 2 specs; [East] Ecuador Macas No. 166; Zuña on Huilca MUSEUM PARIS de Macas Cap D'espanay Mars; Teneb_base 23189 to 23190; NHMB • 2 specs; same location as for preceding; Teneb_base 23179 to 23180; NHMB • 1 spec.; Panelos(?) [illegible]; Teneb_base 23185; NHMB • 8 specs; Napo Prov. Archidona; Rio Hollin; 21 Dec. 2001; R. Diaz, P.Bravo and E. Vasquez leg.; Teneb_base 23234 to 23241; TAMU. GUYANA • 1 spec.; Reg 8, Potaro River, 5°18' N, 59°54.635' W [WGS84]; 5 Jun. 2012; R.A. Koch leg.; Teneb_base 23242; PERC. PERU • 1 spec.; Upper Rio Huallaga; 1 Jan. 1926; Teneb_base 23230; AMNH • 1 spec.; same data as for preceding; Oct. 1929; Teneb_base 23231; AMNH • 1 spec.; Rio Huallaga; Oct. 1930; Teneb_base 23232; AMNH • 1 spec.; same location as for preceding; Teneb_base 23243; AMNH • 3 specs; Kaieteur, BG.; 6 Aug. 1911; Teneb_base 23217 to 23219; AMNH • 2 specs; Marcapata; Teneb_base 23187 to 23188;NHMB•1spec.;CuscoDept.VillaCarmenfieldstation,1.2kmNofcafeteria;12.87753°S, 71.40153° W; 525–1000 m a.s.l.; 27 May 2011; D.J. Bennett leg.; hand collecting; PER-11-DJB-036.; Teneb_base 18774; SEMC. VENEZUELA • 1 spec.; Mt Duida, 1158 m a.s.l., Amazonas; Feb. 1968; J. Rivero; MCZ-Ent 00753334; Teneb_base 23244; MCZ • 1 spec.; La Escalera Bolivar, on rotten log near a stream; 1000 m a.s.l.; 22 Mar. 1968; J.A.Rivero leg.; MCZ-ENT 000753333; Teneb_base 23216; MCZ. Redescription MeasureMents. Length 27–36 mm, width 10–12 mm, specimens measured (n = 20). Head. Medial edge of mandibles without tooth, inner margin without crenulation. Galea with cluster of bristles situated near midpoint. Mentum weakly sculptured, apex with medial notch; lateral apices produced. Clypeal apex not dilated. Supraorbital tubercles absent. Clypeal horn with wide base, arced (inlateralview);dorsalsurfaceflattened,endingbluntlyattip.
European Journal of Taxonomy 1012: 49–80 (2025) 66 Status Whiletheoriginaldescriptiondoesnotprovideaspecificnumberofspecimensexamined;itdoes indicate that there were multiple individuals by providing a size range. Therefore, we designate here a lectotype from the presumed syntype series (see above for label data). Distribution South America: Bolivia, Chile, Ecuador, Guyana, Peru, Venezuela (Fig. 7). Phrenapates educator Gebien 1910 Figs 3F, H, 4E–F, 6, 10A, 13 Phrenapates educator Gebien, 1910: 504. Diagnosis The three species P. erratus Rincon,Lumen&Kamińskinom.nov., P. educator, and P. latreillei are noticeablysmallerthantheotherfivespecies(21–28mminlengthvs26–39mminlargerspecies). Fig. 13. Phrenapates educator Gebien 1910, lectotype, ♂ (Teneb_base 23175, NMHB), dorsal (top) and lateral (bottom) views.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 67 Phrenapates erratus was only represented here by the lectotype (Fig. 14), and no additional material was found during this revision. Gebien (1911) noted that P. erratus is very similar to P. educator. Characters he listed were: P. educator displays a strongly concave lower lip (clypeal apex) vs P. erratus, mandibles that are more deeply hollowed out, and the slightly weaker humeral angles of the elytra that show the epipleural keel (which is covered in P. erratus). Upon comparison of P. erratus vs P. educator, many of thesecharacterswereeitherdifficulttodelineate,orfoundtoincludesomevariationwithinP. educator. Phrenapates educator is best separated from P. erratus by the distinct carina near the attachment point of the mandible, P. erratus has, instead, a weak carinate bump. The dorsal depression on the mandible is deeply punctate without setae, while the mandible of P. erratus is semi-rugose and has setae. The mentum, though variable, tends to be arcuate in P. erratus and sinuate in P. educator. Type material Lectotype (designated here) ECUADOR • ♂;(greenlabel)“W.Ecuador/Pucay300m/(Ohaus)XI.05.//Phrenapates / educator/ X Geb // (red label) Type / No. 163 // Sammlung / H Gebien // (blue label) Tenebrionid Base / Aaron D. Smith / Catalog # 23175”; Teneb_base 23175; NHMB. Other material examined COLOMBIA • 1 spec.; Aguaral; Teneb_base 23183; NHMB. ECUADOR • 1 spec.; St Domingo, Colorados; Jun. 1981; G. Onoré leg.; Teneb_base 23245; CASC • 4 specs; Balzapampa Route de Quita Prov. de Bolivar; Sep.–Oct. 1893; M.de Mathan leg.; Teneb_base 23285 to 23288; MNHN • 2 spec.; 1922; Museum Paris Quito Cap D'Espinay; Teneb_base 23289 to 23290; MNHN • 1 spec.; W. Ecuador Pucay; 300 m a.s.l.; Nov. 1905; Ohaus leg.; Teneb_base 23169; NHMB. Redescription MeasureMents. Length 21–26 mm, width 7–8 mm, specimens measured (n = 6). Head. Medial edge of mandibles without tooth, inner margin without crenulation. Galea with cluster of bristles situated toward apex. Mentum moderately sculptured, apex with medial notch; lateral apices produced. Supraorbital tubercles absent. Clypeal horn weakly arced (in lateral view); ending bluntly at tip. Status Whiletheoriginaldescriptiondoesnotprovideaspecificnumberofspecimensexamined;itdoesindicate that there were multiple individuals by providing a size range (Gebien 1910). Therefore, we designate here a lectotype from the presumed syntype series (see above for label data). Distribution South America: Ecuador, Columbia (Fig. 6). Phrenapates erratus Rincon,Lumen&Kamińskinom.nov. Figs 2D, 3C, 6, 10C, 14 Phrenapates latreillei Gebien, 1910: 504 [junior primary homonym of Phrenapates latreillei Lacordaire, 1859: 315].
European Journal of Taxonomy 1012: 49–80 (2025) 68 Diagnosis For separation of Phrenapates erratus Rincon,Lumen&Kamińskinom.nov.fromlargerspecies and from P. latreillei see the diagnoses for P. latreillei and P. educator, respectively. Morphologically, P. erratus canbedifficulttodifferentiatefromP. educator. Phrenapates erratus is best separated from P. educator in the weak carinate bump near the attachment point of mandible, P. educator has a distinct carina in the proximal end. The dorsal depression on mandible is rugose and has setae in P. erratus while in P. educator the dorsal depression is deeply punctate and does not have setae. The mentum, though variable, tends to be arcuate in P. erratus while it is sinuate in P. educator. Etymology Named in reference to remedying or editing previously authored taxon names. Type material Lectotype (designated here) UNKNOWNLOCATION•♂;(greenlabel)“Nov.Granada//Phrenapates / Latreillei / Geb // (red label) Type / No. 166 // Sammlung / H Gebien // (blue label) Tenebrionid Base / Aaron D. Smith / Catalog # 23168”; Teneb_base 23168; NHMB. Fig. 14. Phrenapates erratus Rincon,Lumen&Kamińskinom.nov.,lectotype,♂ (Teneb_base 23168, NHMB), dorsal (top) and lateral (bottom) views.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 69 Redescription MeasureMents. Length 23 mm, width 8 mm (n = 1). Head. Medial edge of mandibles without tooth, inner margin without crenulation. Galea with cluster of bristles situated toward apex. Mentum strongly sculptured, apex with medial notch; lateral apices produced. Clypeal apex dilated. Supraorbital tubercles absent. Clypeal horn bent weakly downward (in lateral view); ending bluntly at tip. Status Gebien’sfirsttreatment(1910)ofPhrenapates presents P. latreillei as a new species in preparation for a subsequenttreatment.In1911,Gebienattemptedtostabilizethenomenclaturebyofficiallyintroducing the name Phrenapates latreillei forspecimensavailabletohimthatwere,inhisopinion,fittingthe speciesconcept.Gebien’s(1910,1911)treatmentsappeartobebasedonthepresumptionthat“in litt.” attributions to Dejean (i.e. Gebien 1911, Gemminger and Harold 1870) and Lacordaire’s (1859) species description were invalid. His initial description was based on more than a single specimen (inferred from size range provided) from New Granada (Gebien 1910); however, in 1911 when providing a more detailed redescription of this species, he stated that only a single specimen was available to him, which is here designated as the lectotype to stabilize the concept of the species (see label data above).Furthermore,inthesamepaper,Gebien(1911)alsoattributedthisspeciestoDejean(“inlitt.”), introducing organizational chaos in the nomenclature. Reevaluation of the literature has revealed that Lacordaire’s (1859) description of Phrenapates latreillei does, in fact, meet the requirements of the ICZN (1999) and should be treated as valid. As such, Gebien’s Phrenapates latreillei should be treated as a junior primary homonym of Lacordaire’s (1859) name. A replacement name, Phrenapates erratus Rincon,Lumen&Kamiński nom. nov., is hereby introduced for Phrenapates latreillei Gebien, 1910. Finally,verificationofhistoricaltypematerialshowsthatGebien’s(1910)conceptofP. latreillei does not match the concept presented by Lacordaire (1859). As such, P. erratusRincon,Lumen&Kamiński nom. nov. is hereby treated as a separate species. The investigation conducted here supports a close relation between this species and Phrenapates educator; however, P. erratus remains diagnosable via the weak carina near the proximal end of the mandible and the rugosity of the mandible and the presence of setae. More material is needed to fully test the status of Phrenapates erratus. Finally, we designate aspecimenidentifiedasP. latreillei by Gebien in NHMB as the lectotype for Phrenapates erratus (see Type Material of P. erratus above). Distribution South America: the label information cannot be georeferenced as it refers to a large geographic region (New Granada), which includes areas of Columbia, Ecuador, Panama, and Venezuela. Phrenapates fortunaensis Rincon & Smith sp. nov. urn:lsid:zoobank.org:act:92CCAB44-7C22-4F30-A36F-79139C448612 Figs 2G, 3E, 6–7, 15 Diagnosis Phrenapates fortunaensis Rincon & Smith sp. nov. can be separated from all other known congeners based on the following characters: supraorbital tubercles present, separating this species from P. dux, P. latreillei, P. educator, and P. erratus Rincon,Lumen&Kamińskinom.nov.Thesupraorbitaltubercles are wider than tall, similar to P. bennettii, but different than in P. gilloglyi Rincon & Smith sp. nov. Mandibles with crenulations on inside margin, as well as with an extra tooth as in P. bennettii, which
European Journal of Taxonomy 1012: 49–80 (2025) 70 differs from the smooth inner margin of P. gilloglyi. The mentum lacks an apical medial notch, instead possessing a small projection, giving the appearance of a trilobate mentum (Fig. 2G). Etymology Species named after the Fortuna Mountains where the specimens were found. Type material Holotype PANAMA • Chiriqui Prov., Reserva Fortuna, 3rdNov.Trail;8°45.468′N,82°15.671′W;1234–1280m a.s.l.; 1 Sep. 2008; A.R. Gillogly leg.; Teneb_base 23482; PERC. Other material examined PANAMA • 1 spec.; Boca d. Toro Prov., 3 km NW of Fortuna Hwy border station; 900 m a.s.l.; 1 Aug. 1997; A.R. Gillogly leg.; Teneb_base 23331; AAAG • 2 specs; Bocas del Toro Prov., La Culebra Trail, 5 km N of Boquete; 1500 m a.s.l.; 16 Jul. 1995; A.R. Gillogly leg.; Teneb_base 23468 to 23469; AAAG • 1 spec.; Chiriqui Prov. Reserva la Fortuna Cont. Div. Trail; 1300 m a.s.l.; 8°47' N, 82°13' W; 18 Jan. 1992; A.R. Gillogly leg.; Teneb_base 23324; AAAG • 4 specs; Chiriqui Prov, Reserva Fortuna, 3 rd Nov. Trail; 1234–1280 m a.s.l.; 8°45.468' N, 82°15.671 W; 1 Sep. 2008; A.R. Gillogly leg.; Teneb_base 23482 to 23485; AAAG • 3 specs; Chiriqui Prov. Rsva la Fortuna Cont. Div.Trail; 1200 m a.s.l.; 27 Apr. 1996; A.R. Gillogly leg.; Teneb_base 23323, 23466 to 23467; AAAG • 10 specs; Chiriqui Prov. Reserva Fig. 15. Phrenapates fortunaensis Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23482, PERC), dorsal (top) and lateral (bottom) views.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 71 Fortuna, Fortuna Hwy, N of Dam; 1151 m a.s.l.; 8°24.840' N, 82°14.562' W; 30 Aug. 2008; Gillogly and Smith leg.; Teneb_base 23472–23481; AAAG • 2 specs; Chiriqui Prov. Reserva la Fortuna, powerline cut; 1145 m a.s.l.; 8°44.80' N, 82°13.90' W; 31 Jul. 2008; A.Gillogly and A. Smith leg.; Teneb_base 23486 to 23487; AAAG • 5 specs; Chiriqui Prov. Rsva Fortuna, 1.7 km N of Dam; 1152 m a.s.l.; 8°44.829' N, 82°14.572' W; 25 Apr. 2009; A. Gillogly and D. Ward Jr. leg.; Teneb_base 23317 to 23321; AAAG • 2 specs; Chiriqui Prov. Rsva Fortuna, 3 rd Nov. Trail; 1527 m a.s.l.; 8°45.768' N, 82°15.358' W; 28 Apr. 2009; A. Gillogly and D. Ward leg.; Teneb_base 23315 to 23316; AAAG • 2 specs; Chiriqui Prov. Rsva Fortuna, Samudio Trl.; 1090 m a.s.l.; 8°44.1' N, 82°14.8' W; 27 Apr. 2009; A.Gillogly and D.Ward Jr. leg.; Teneb_base 23470 to 23471; AAAG • 3 specs; Chiriqui Reserva la Fortuna, Est. Biologica; 1200 m a.s.l.; 8°43'18" N, 82°14'17" W; 8 Jul. 1999; A.R. Gillogly leg.; Teneb_base 23330, 23464 to 23465; AAAG • 1 spec.; Chiriqui, 3rd Nov. Trail, on rotten logs; 8°45.47' N, 82°15.67' W; 1227 m a.s.l.; 1 Sep. 2008; A.D. Smith leg.; Teneb_base 18775; ADSC • 6 specs; Chiriqui, Powerline cut; 1 km NW of Charchita; 8°24.840' N, 82°14.562' W; 1151 m a.s.l.; 30 Aug. 2008; A.D. Smith leg.; Teneb_base 18766, 18769 to 18773; ADSC • 2 specs; Chiriqui, Powerline cut, N of Lago, Fortuna: 1163 m a.s.l.; 8°44.75' N, 82°13.94' W; 31 Aug. 2008; A.D. Smith leg.; Teneb_base 18777 to 18778; ADSC • 1 spec.; Chiriqui Fortuna; 82°15' W, 8°44' N; 20 May 1978; O'Briens and Marshall leg.; Teneb_base 22814; CASC • 3 specs; Chiriqui Reserva La Fortuna, Hydrographic sta. trail; 28 May 1993; E.G. Riley; Teneb_ base 22811–22813; TAMU. Description MeasureMents. Length 26–33 mm, width 9–11 mm, specimens measured (n = 20). Head. Medial edge of mandibles with tooth, inner margin crenulated. Galea with cluster of bristles situated near midpoint. Mentum moderately sculptured, apex without medial notch; lateral apices rounded. Clypeal apex not dilated. Supraorbital tubercles present; shorter than wide. Clypeal horn very short, weakly arced (in lateral view) and weakly tapered to blunt point. Distribution Central America: Panama. Phrenapates gilloglyi Rincon & Smith sp. nov. urn:lsid:zoobank.org:act:76CB49A2-518F-406D-B9CE-F161967E351A Figs 2B, 3F, 6–7, 8E–F, 16 Diagnosis Phrenapates gilloglyi Rincon & Smith sp. nov. can be separated from all other known congeners based on the following characters: supraorbital tubercles present, separating this species from P. dux, P. latreillei, P. educator, and P. erratus Rincon,Lumen&Kamińskinom.nov.Supraorbitaltubercleswithaheight greater than or equal to the width, separating them from P. bennettii and P. fortunaensis Rincon & Smith sp. nov. Clypeal horn ending in three points, unlike the smoothly tapered horn in P. ohausi. Mandibles with no crenulations or teeth on inside margin, which differs from P. bennettii and P. fortunaensis. Mentum with apical notch, unlike in P. fortunaensis or P. bennettii. Etymology The species is named after the passalid taxonomist Dr Alan Gillogly for his donation of specimens and valuable insight.
European Journal of Taxonomy 1012: 49–80 (2025) 72 Type material Holotype PANAMA•♂;“Panama,CocleProv.,3kmNofElValle,CerroGaital;900ma.s.l.;21Jul.1999;A. Gilloglyleg.;8°37′35″N,80°6′53″W;Teneb_base23367;TenebrionidBase/AaronD.Smith/Catalog # 23367”; Teneb_base 23367; PERC. Other material examined PANAMA • 3 specs; Panama Prov. Cerro Jefe; 894 m a.s.l.; 9°13.339' N, 79°22.257' W; 18 Aug. 2008; Gillogly and Smith leg.; Teneb_base 23364 to 23366; AAAG • 2 specs; Cocle Prov., 3 km N of El Valle Cerro Gaital; 900 m a.s.l.; 8°37'35" N 80°6'53" W; 21 Jul. 1999; A. Gillogly leg.; Teneb_base 23367 to 23368; AAAG • 1 spec.; Cocle Prov. El Valle, Cerro Gaital; 850 m a.s.l.; Jun. 2008; Daniel Curoe leg.; Teneb_base 23369; AAAG • 1 spec.; Bocas d. Toro Prov. Tank Hill, 5 km S of Chiriqui Grande; 750 m a.s.l.; 1 Aug. 1997; A.R. Gillogly leg.; Teneb_base 23370; AAAG • 1 spec.; Cerro Jefe road to Altos de Pacora; 894 m a.s.l.; 9°13.34' N, 79°22.26' W; 18 Aug. 2008; A.D. Smith leg.; Teneb_base 18780; ADSC • 1 spec.; Cerro Jefe summit, Panama Prov.; 8–10 May 1985; F.T. Hovore leg.; Teneb_base 22950; CASC • 3 specs; Cerro Jefe; 700–800 m a.s.l.; 26 Oct 1969; B. Malkin leg.; Teneb_base 23356 to 23358; FMNH • 1 spec.; Rio Changena Bocas del Toro R.PAN; Sep. 1961; G.B. Fairchild leg.; Teneb_base 23322; MCZ. Fig. 16. Phrenapates gilloglyi Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23367, PERC), dorsal (top) and lateral (bottom) views.
RINCON A. et al., Revision of the darkling beetle genus Phrenapates (Tenebrionidae: Phrenapatinae) 73 Description MeasureMents. Length 26–33 mm, width 9–11 mm, specimens measured (n = 12). Head. Medial edge of mandibles without tooth, inner margin without crenulation. Galea with cluster of bristles situated near midpoint. Mentum weakly sculptured, apex with medial notch; lateral apices rounded. Supraorbital tubercles present; height greater than width. Clypeal horn straight to weakly arced (in lateral view); ending in three, raised blunt points. Distribution Central America: Panama (Fig. 7). Phrenapates latreillei Lacordaire, 1859 Figs 2E, 3G, 4G–H, 6–7, 9A, 17 Phrenapates latreillei Lacordaire, 1859: 315. Phrenapates mandibularis Gebien, 1910: 504. Syn. nov. Diagnosis Phrenapates latreillei has no supraorbital tubercles, separating it from P. bennettii, P. ohausi, P. gilloglyi Rincon & Smith sp. nov. and P. fortunaensis Rincon & Smith sp. nov. The horn of P. latreillei (Fig. 3G) differs from the unique form of P. dux (Fig. 3D). Phrenapates latreillei (as well as P. educator) can usually be separated from P. dux (length 27–36 mm) as sizes range from 18–28 mm. The closest species morphologically to P. latreillei is P. educator. Phrenapates latreillei has a horn with a stronger arc than P. educator (Fig. 3G–H, respectively). Additionally, there is a tooth on the mandible which is absent in P. educator. The mentum has stronger paramedial ridges and a deeper medial excavation in P. latreillei vs the less sculptured mentum in P. educator. Though not always consistent, the apex of the labrum is often more convex in P. latreillei, whereas it tends to be straight/slightly concave in P. educator. Type material Holotype [Phrenapates mandibularis syn. nov.] COLOMBIA•♂;(greenlabel)“00;//Phrenapates / mandibularis / X Geb // (red label) Type / No. 165 // Sammlung / H Gebien // (blue label) Tenebrionid Base / Aaron D. Smith / Catalog # 23167”; Teneb_base 23167; NHMB. Lectotype (designated here) COLOMBIA•♂;(greenrectanglelabel)“Phrenapates / Latreillei Dj / Columbia [N J Les] // (green circle label) [unintelligible] / Latreillei Dej / Columb / [unintelligible]”; MNHN. Other material examined BRAZIL • 1 spec.; Roraima; Whiteley; 6000 m a.s.l.; Teneb_base 23279; MNHN. COLOMBIA • 1 spec; s. loc.; Teneb_base 23278; MNHN • 2 specs; s. loc.; Teneb_base 23276 to 23277; repository unknown. PANAMA • 36 specs; Darien Prov. Cana, Pirre Trail; 1560 m a.s.l.; 5 Jun. 1996; A.R Gillogly leg.; Teneb_base 23246, 22740 to 22746, 22764 to 22780, 22798 to 22808; AAAG • 10 specs; Darien Prov. Cana, Pirre Ridge; 1650 m a.s.l.; 7°45.825' N, 77° 43.325' W; 5 May 2008; A.R Gillogly and M. Seid leg.; Teneb_base 23247, 22730 to 22738; AAAG • 1 spec.; Bocas d. Toro Prov., 3 km N of Cont. Div. Trail, Fortuna Hwy; 950 m a.s.l.; 13 Jul. 1996; A.R. Gillogly leg..; Teneb_base 22739; AAAG.
European Journal of Taxonomy 1012: 49–80 (2025) 80 Schawaller W. & Bouchard P. 2019. New taxa of Phrenapatinae (Coleoptera: Tenebrionidae) from Southern Africa, with new distributional data and a checklist of the Afrotropical fauna. Annales Zoologici 69 (1): 191–200. https://doi.org/10.3161/00034541ANZ2019.69.1.014 Schuster J. & Schuster L. 1985. Social behavior in passalid beetles (Coleoptera: Passalidae): cooperative brood care. Florida Entomologist 68: 266–272. https://doi.org/10.2307/3494359 Schuster J. & Schuster L. 1997. The evolution of social behavior in Passalidae (Coleoptera). In: Choe J.C. & Crespi B. (eds) The Evolution of Social Behavior in Insects and Arachnids: 260–269. Cambridge University Press, Cambridge, USA. https://doi.org/10.1017/CBO9780511721953.013 Solier A.J. 1834. Essai d’une division des Coléoptères Hétéromères, et d’une monographie de la famille des Collaptèrides. Annales de la Société entomologique de France 3: 479–636. Still C.J., Foster P.N. & Schneider S.H. 1999. Simulating the effects of climate change on tropical montane cloud forests. Nature 398: 608–610. https://doi.org/10.1038/19293 Tschinkel W.R. & Doyen J.T. 1980. Comparative anatomy of the defensive glands, ovipositors and female genital tubes of tenebrionid beetles (Coleoptera). International Journal of Insect Morphology and Embryology 9 (5–6): 321–368. https://doi.org/10.1016/0020-7322(80)90009-4 WattJ.C.1974.ArevisedsubfamilyclassificationofTenebrionidae(Coleoptera). New Zealand Journal of Zoology 1 (4): 381–452. https://doi.org/10.1080/03014223.1974.9517846 Wicknick J.A & Miskelly S.A. 2009. Behavioral interactions between non-cohabiting bess beetles, Odontotaenius disjunctus (Illiger) (Coleoptera: Passalidae). The Coleopterists Bulletin 63 (1): 108–116. https://doi.org/10.1649/0010-065X-63.1.108 Printed versions of all papers are deposited in the libraries of four of the institutes that are members of the EJTconsortium:MuséumnationaldʼHistoirenaturelle,Paris,France;MeiseBotanicGarden,Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences, Brussels, Belgium. The other members of the consortium are: Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the Czech Republic, Prague, Czech Republic; The Steinhardt Museum of Natural History, Tel Aviv, Israël.