Phrenapates 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 : 52-59

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 meso- and 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.

Redescription

MEASUREMENTS. Length 18–39 mm, width 6–13 mm, specimens measured (n = 105). Color nitidus dark brown to black. Body sub-cylindrical.

HEAD. 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

Member of the subfamily Phrenapatinae in tribe Phrenapatini , which is unified by the mesocoxae lacking 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 & Kamiński nom. 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.

Key to the species of Phrenapates Gray, 1832

1. Two supraocular tubercles on the head ( Fig. 8A–F ) ......................................................................... 2

– Lacking supraocular tubercles ( Fig. 8G–H ) .................................................................................... 5

2. Clypeal horn tapered to a point ( Fig. 3A ); mandibles without crenulation or tooth on the inner margin (as in Fig. 9C ); supraorbital tubercles height subequal to width ( Fig. 8A–B ), mentum with rounded front angles ( Fig. 2H ) .................................................................................... P. ohausi Gebien, 1910

– Clypeal horn with end either blunted or with three points ( Fig. 3B, F ); mandibles with or without crenulations or tooth on the inside edge; supraorbital tubercles taller than wide or shorter than wide ( Fig. 8C–F ), mentum with lateral apices produced (as in Fig. 2A, C–D ) ........................................ 3

3. Clypeal horn ending in three points ( Fig. 3F ); supraorbital tubercles taller than wide; inner mandible smooth, without crenulated teeth (as in Fig. 9C ) ....................... P. gilloglyi Rincon & Smith sp. nov.

– Clypeal horn with single, blunt ending ( Fig. 3B, E ); supraorbital tubercles not as tall as wide ( Fig. 8C–D ); mandible with crenulations on inner margin ( Fig. 9B ) ............................................... 4

4. Lateral apices of mentum rounded ( Fig. 2G ); clypeal horn short (not or barely arced) ( Fig. 3E ); mandibles with tooth on inner margin (as in Fig. 9A ) ....... P. fortunaensis Rincon & Smith sp. nov.

– Lateral apices of mentum produced ( Fig. 2A ); clypeal horn arced (on rare occasions short) ( Fig. 3B ); mandibles without tooth and with crenulations on inner margin ( Fig. 9B ) ... P. bennettii Gray, 1832

5. Clypeal horn with slightly enlarged base laterally, tapering in center, with broad and flattened top ( Fig. 3D ); never with tooth on mandible ( Fig. 9C ); large species ( 27–32 mm long) ......................... ............................................................................................................................. P. dux Gebien, 1910

– Clypeal horn not as above; with or without tooth on mandible (as in Fig. 9A, C ); generally smaller species (< 28 mm long) ..................................................................................................................... 6

6. Mandibles with tooth on inner margin (as in Fig. 9A ); clypeal horn curved in strong, circular arc ( Fig. 3G ); small bundle of bristles placed near midpoint of galea ....... P. latreillei Lacordaire, 1859

– Mandibles without tooth on inner margin ( Fig. 9C ); clypeal horn weakly or not at all curved downward ( Fig. 3C, H ); small bundle of bristles placed nearer to apex of galea ( Fig. 10A, C ) ...... 7

7. Carinate ridge near attachment point of mandible, dorsal depression semi-rugose and with setae, mentum sinuate on sides, ratio of distance from clump of bristles on galea to tip and to base is 1:2 ( Fig. 10A ), Ecuador.................................................................................... P. educator Gebien, 1910

– Carinate ridge near attachment point of mandible weaker, dorsal depression not semi-rugose but punctate and without setae, mentum arctuate on sides, ratio of distance from clump of bristles on galea to tip and to base is less than 1:2 ( Fig. 10C ), Columbia ........................................................... .............................................................................. P. erratus Rincon, Lumen & Kamiński nom. nov.

Figures

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. 7. Distribution of the Phrenapates dux Gebien, 1910, Phrenapates latreillei Lacordaire, 1859, Phrenapates gilloglyi Rincon & Smith sp. nov., and Phrenapates fortunaensis Rincon & Smith sp. nov.

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. 10. Position of galea setae cluster in Phrenapates Gray, 1832. A. Near the apex (ratio tip to clump of bristle to tip to end 1:2) as in Phrenapates educator Gebien, 1910, lectotype, ♂ (Teneb_base 23175, NMHB). B. Centered as in Phrenapates ohausi Gebien, 1910, ♂ (Teneb_base 23165, NHMB). C. Galea ratio less than 1:2 in Phrenapates erratus Rincon, Lumen & Kamiński nom. nov., lectotype, ♂ (Teneb_ base 23168, NHMB).