Phrenapates bennettii Gray, 1832

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, pp. 49-80 : 61-63

Phrenapates bennettii Gray, 1832

Figs 1B , 2A , 3B , 4A–B , 5A , 6 , 8C–D , 9B , 11

Phrenapates bennettii Gray, 1832: 91 .

Diagnosis

Phrenapates bennettii 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 & Kamiński nom. nov. The supraorbital tubercles in P. bennettii are shorter than wide ( Fig. 8C–D ), separating them from P. gilloglyi (tubercles are taller than wide) and P. ohausi (tubercles are as tall as wide). Clypeal horn ends in a blunted point, unlike the smooth tapered horn in P. ohausi , short horn in P. fortunaensis , and three-pointed horn in P. gilloglyi . Mandibles have crenulations on the inside margin, which differs from P. gilloglyi and P. ohausi . Phrenapates bennettii have variation in the shape of their mentum but never have a notch like that of P. gilloglyi or P. ohausi ( Fig. 2B, H ); the mentum has produced lateral apices which differs from P. fortunaensis with rounded lateral apices.

Type material

Lectotype (designated here)

COLUMBIA • ♂; “ Phrenapates / Bennettii / Kirby, Columbia // F. Bates / 81.19 // (red label) LECTOTYPE / Phrenapates / bennettii / Gray 1832 // (blue label) Tenebrionid Base / Aaron D. Smith / Catalog # 23170”; Teneb_base 23170; BMNH.

Paralectotypes

The two additional specimens from Kirby’s material should be considered paralectotypes. However, we were unable to locate them during this study.

Other material examined

COLUMBIA • 1 spec.; Bogota; 1924; Teneb_base 23174; NHMB .

COSTA RICA • 2 specs; Puntarenas, Cote Brus, Est Biol. Las Alturas ; 8°57' N, 82°52' W; 26 Mar. 2002; in rotten logs; A. Cline and A. Tishechkin leg.; Teneb_base 22871 to 22872; AAAG • 1 spec.; Provincia de Puntarenas, Canton Coto Brus. Agua Caliente ; Aug. 1986; Nick Zakharoff gift to Cal. Acad. Soi. Coll.; Teneb _base 23263; CASC • 1 spec.; Cartago 15 km NE of Turrialba; 18–21 Apr. 1981; E. Giesbert leg.; Teneb_base 23261; CASC • 1 spec.; Bebedero [Underwood, 94]; Teneb_base 23265; MNHN • 1 spec.; San Carlos [78 Plesiotype 161]; Teneb_base 23186; NHMB • 1 spec.; Puntarenas 8 km S of San Vito; 25 May 1983; Dan Heffern leg.; Teneb_base 23262; TAMU .

ECUADOR • 2 specs; Cachabe. [low c.]; Nov. [96]; Rosenberg leg.; Teneb_base 23254 to 23255; MNHN • 1 spec.; Equateur Loja Abbé Guajon ; Teneb_base 23253; MNHN .

GUATEMALA • 3 specs; Teneb_base 23250 to 23252; MNHN • 3 specs; St Cristobal , Verapaz; Teneb_ base 23156 to 23158; NHMB • 1 spec.; s. loc.; Teneb_base 23172; NHMB .

HONDURAS • 1 spec.; [Coll. Nonfried]; Teneb_base 23256; MNHN • 1 spec.; s. loc.; Teneb_base 23171; NHMB .

NICARAGUA • 18 specs; Jinotega, Cerro Kilambé ; 1000 m a.s.l.; Jul. 2001; col. M.A. Guatemala; Teneb_base 23249, 22901, 22903 to 22916, 22935 to 22936; AAAG • 6 specs; Zelaya, Cerro Saslaya; 500 m a.s.l. ( Bosque ); 7–14 Jul. 1999; B. Hernandez leg.; Teneb_base 22937 to 22942; AAAG • 1 spec.; San Carlos, Rio San Juan, Est. Biol. [Bartola] Selva alta perenfolia ; 30 m a.s.l.; 10°58' N, 84°20' W; 11 Sep. 2000; J. Marquez leg.; en troncos caidos; Teneb_base 22943; AAAG • 1 spec.; Bluefields ; 1–3 Apr. 1949; L.E. Long leg.; Teneb_base 23264; FMNH .

PANAMA • 10 specs; Chiriqui prov., 5 km NE of Santa Clara, Hartmann's ; 1515 m a.s.l.; 8°50.68' N, 82°44.58' W; 23 Aug. 2008; A. Gillogly leg.; Teneb_base 22838 to 22847; AAAG • 1 spec.; 9.7 km W of Volcan Chiriqui prov.; 28 Dec. 1991; 1400 m a.s.l.; A.Gillogly and J.H. White leg.; Teneb_base 22880; AAAG • 2 specs; Cocle Prov., 3 km N of El Yalle, La Mesa ; 930 m a.s.l.; 8°38'13" N, 80°07'28" W; 21 Jul. 1999; A.R. Gillogly leg.; Teneb_base 22848, 22867; AAAG • 2 specs; Cocle Prov., 3 km N of El Valle ; Teneb_base 22868 to 22869; AAAG • 1 spec.; La Mesa , NE side of Cerro Gaital , Cocle Prov.; 850–900 m a.s.l.; 13 Oct. 1991; A.R. Gillogly leg.; Teneb_base 22870; AAAG • 4 specs; Cocle Prov., NE side of La Mesa , Cerro Gaital ; 900 m a.s.l.; 1 Mar. 1992; A.R. Gillogly leg.; Teneb_base 22876 to 22879; AAAG • 1 spec.; 10 km N of El Cope, Cocle Prov.; 750 m a.s.l.; 21 Dec. 1991; Teneb_base 22881; AAAG • 22 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 22944 to 22948, 23248, 23332 to 23347; AAAG • 25 specs; Darien Prov. Cana, Pirre Trail ; 1560 m a.s.l.; 5 Jun. 1996; A.R Gillogly leg.; Teneb_base 22969 to 22984, 23348 to 23355, 22902; AAAG • 6 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 22832 to 22837; AAAG • 2 specs; Veraguas Prov., 8 km W of Santa Fe Cerro Tute; 950 m a.s.l.; 8°30'26" N, 81°6'49" W; 24 Jul. 1999; A.Gillogly leg.; Teneb_base 22873 to 22874; AAAG • 1 spec.; Veraguas Prov. 4km N of Alto de Piedra, NW Sta Fe .; 900 m a.s.l.; 10 Apr. 1993; A.R. Gillogly leg.; Teneb_base 22875; AAAG • 1 spec.; 4.8 km NNE of Santa Fe Veraguas, Panama Prov.; 700 m a.s.l.; 8 Dec. 1991; Teneb_base 22882; AAAG • 1 spec.; 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 18779; ADSC • 1 spec.; Barro Colorado , C.Z.; 3 Feb. 1936; W.J. Gertsch leg.; Teneb_base 23300; AMNH • 1 spec.; Barriles , Chiriqui; 31 Jan. 1931; Teneb_base 23302; CASC • 1 spec.; Potrerilles ; 25 Jan. 1935; Teneb _base 23303; CASC • 1 spec.; Bocas Del Toro, 10 km NE of Fortuna Dam; 1036 m a.s.l.; 23–26 May 1984; E. Giesbert leg.; Teneb_base 23301; CASC • 1 spec.; Chiriqui; Champion ex. Gordon and Salvin leg.; Teneb_base 23299; MCZ • 1 spec.; Chiriqui; Teneb_base 23304; MNHN • 10 specs, Museum Paris ex coll. E. Oberthur; V. de Chiriqui m de Mathan ; 1901; Teneb_base 23305 to 23314; MNHN • 1 spec.; Cocle Prov. Cerro, Gaitai ; 1000 m a.s.l.; 8°36' N, 80°10' W; 11 Nov. 1992; P. Jolivet leg.; Teneb_base 23260; MNHN • 4 specs; Chiriqui; Teneb_base 23162 to 23164, 23178; NHMB • 1 spec.; Chiriqui; Teneb_base 23173; NHMB • 1 spec.; [Chiriqui 59]; Teneb_base 23177; NHMB .

UNKNOWN • 3 specs; patria; Teneb_base 23159 to 23161; NHMB.

Redescription

MEASUREMENTS. Length 26–39 mm, width 9–13 mm, specimens measured (n = 20).

HEAD. Medial edge of mandibles without tooth, inner margin crenulated. Galea with cluster of bristles situated near midpoint. Mentum weakly sculptured, apex without medial notch; lateral apices produced. Clypeal apex not dilated. Supraorbital tubercles present; shorter than wide. Clypeal horn arced (in lateral view); ending bluntly at tip.

Status

Gray in Griffith and Pidgeon (1832) did not designate a holotype from the type series of Kirby’s three specimens from Choco, Columbia. Therefore, a lectotype designation is needed to stabilize the taxonomic status of the species and genus. There is one specimen from Kirby’s collection held by the Natural History Museum in London (BMNH) with the correct locality, which is hereby designated as the lectotype.

Distribution

Central America: Costa Rica, Guatemala, Honduras, Nicaragua, Panama; South America: Colombia, Ecuador ( Fig. 6 ).

Figures

Fig. 1. Convergent morphology of Passalidae Leach, 1815, and Phrenapates Gray, 1832. A. Passalidae Leach, 1815 (MIZPAN). B. Phrenapates bennettii Gray, 1832 (MIZPAN).

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_base 23168, NHMB). E. Phrenapates mandibularis Gebien 1910 [Phrenapates latreillei Lacordaire], holotype, ♂ (Teneb_base 23167, NHMB). F. Phrenapates educator Gebien, 1910, lectotype, ♂ (Teneb_base 23175, NHMB). G. Phrenapates fortunaensis Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23482, PERC). H. Phrenapates ohausi Gebien, 1910, lectotype, ♂ (Teneb_base 22810, BMNH).

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 & Kamiński nom. nov., lectotype, ♂ (Teneb_base 23168, NHMB). D. Phrenapates dux Gebien, 1910 (Teneb_base 23215, AAAG). E. Phrenapates fortunaensis Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23482, PERC). F. Phrenapates gilloglyi Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23367, PERC). G. Phrenapates latreillei Lacordaire, 1859, ♂ (Teneb_base 22740, AAAG). H. Phrenapates educator Gebien, 1910 (Teneb_base 23286, MNHM)

Fig. 4. Phrenapates Gray, 1832, adeagi, dorsal and lateral views. A–B. Phrenapates bennettii Gray, 1832, ♂ (Teneb_base 23186, 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_base 23167, NHMB). I–J. Phrenapates ohausi Gebien, 1910, ♂ (Teneb_base 23165, NHMB).

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.

Fig. 8. Supraorbital tubercles of Phrenapates Gray, 1832. A–B. Phrenapates ohausi Gebien, 1910, (Teneb_base 23212, CASC). C–D. Phrenapates bennettii Gray, 1832, (Teneb_base 22972, AAAG). E–F. Phrenapates gilloglyi Rincon & Smith sp. nov., holotype, ♂ (Teneb_base 23367, PERC). G–H. Absence of supraorbital tubercles in Phrenapates dux Gebien, 1910, (Teneb_base 23215, AAAG).

Fig. 9. Mandible crenulations and teeth in Phrenapates Gray, 1832. A. Tooth, no crenulations in Phrenapates mandibularis Gebien 1910 [Phrenapates latreillei], holotype, ♂ (Teneb_base 23167, NHMB). B. Crenulations, no tooth, in Phrenapates bennettii Gray, 1832, ♂ (Teneb_base 23186, NHMB). C. No crenulations or tooth in Phrenapates dux Gebien, 1910 (Teneb_base 23215, AAAG)

Fig. 11. Phrenapates bennettii Gray, 1832, lectotype, ♂ (Teneb_base 23170, BMNH), dorsal (top) and lateral (bottom) views.