Phanaeus pyrois Bates, 1887: 58 Phanaeus pyrois Gillet 1911: 85 Olsoufieff 1924: 37 Edmonds 1972: 830 Howden & Young 1981: 134 Krajcik 2006: 152 Price 2007: 17 Solís & Kohlmann 2012: 1 Edmonds & Zídek 2012: 1 Moctezuma & Halffter 2017: 55 Kohlmann et al . 2018: 69 Chamorro et al . 2019: 220 Phanaeus ( Notiophanaeus ) pyrois Edmonds 1994: 2 Arnaud 2002b: 96 Edmonds & Zídek 2012: 3 Phanaeus ( Notiophanaeus ) pyrois pyrois Arnaud 2002b: 96 Phanaeus pyrois pyrois Arnaud 2002b: 97 Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species Moctezuma, Victor Halffter, Gonzalo European Journal of Taxonomy 2021 2021-04-28 747 1 71  Chamorro et al . 2019: 220  4FSJX Bates, 1887 Bates, Det. W. D. Edmonds 1887 [620,968,264,291] Insecta Scarabaeidae Phanaeus Animalia Coleoptera 41 42 Arthropoda species pyrois   Figs 1N, 2J, 12, 16, 18O, 19O      Phanaeus pyrois Bates, 1887: 58, pl. 2, table 3, figs 22–23 (in part).    Phanaeus pyrois– Nevinson 1982: 6 (in part). —  Gillet 1911: 85(in part). —  Olsoufieff 1924: 37, 93, 152 (in part). — Blackwelder 1944–1957: 210 (in part). —  Edmonds 1972: 830(in part); 1979: 103 (in part); 1994: 3, 5, 8–9, 39, 44–46, 103 (in part). —  Howden & Young 1981: 134, 136 (in part). —  Krajcik 2006: 152(in part). —  Price 2007: 17, figs 52–53, 54 (in part); 2009: 145 (in part). —  Solís & Kohlmann 2012: 1, 8–10, 31, fig. 1 (in part). —  Edmonds & Zídek 2012: 1, 5–6, 8, 13 (in part). —  Moctezuma & Halffter 2017: 55(in part). — Moctezuma et al. 2017: 114, 130 (in part). —  Kohlmann et al. 2018: 69, 78–79, 83, 88, 89, fig. 8a, d (in part). —  Chamorro et al. 2019: 220(in part).    Phanaeus( Notiophanaeus) pyrois–  Edmonds 1994: 2, 8, 41, 44, figs 210, 214–215, 221 (in part). —  Arnaud 2002b: 96(in part). —  Edmonds & Zídek 2012: 3, 13, figs 138, 142–143, 156–159 (in part).    Phanaeus( Notiophanaeus) pyrois pyrois–  Arnaud 2002b: 96(in part).    Phanaeus pyrois pyrois–  Arnaud 2002b: 97(in part).    Diagnosis Easily diagnosed species by the pronotum bright metallic red ( Figs 12A, D), green ( Fig. 2J) or dark metallic blue (12B), with elytral striae not strongly impressed basally ( Fig. 12). The rest of the green/ blue species of the  P. endymionspecies group are recognized by the elytral striae strongly impressed basally as a distinct fossa. A black dorsal colour is never found in  P. pyroisspecimens. Minor males of  P. panamensis sp. nov.and red  P. pyroismay be strongly mimetic, but easily separated by the endophallite copulatrix ( Fig. 1).    Type material   Lectotype(studied from photographs, 1 ♂)   NICARAGUA–  Chontales• ♂, Edmonds 1994: 45( Fig. 12D); “NHMUK 013678267/ B. C. A. p. 58, sp.8. / LECTO-TYPE/  Phanaeus pyroisBates.LECTOTYPE ♂P. ARNAUD DET 1980/T. Belt /Type /  P. pyrois♂/ Sp. figured”; NHMUK 013678267; BMNH.  Non-type material revised( 7 ♂♂, 4 ♀♀)   NICARAGUA–  Granada• 1 ♂; “ Volcán Mombacho. Bosque Seco.  30-VI-98. JM. Mars”; TAMU•  1 ♂; “ Volcán Mombacho. Santa Ana.  21-V-98. Malaise. JM. Mars”; VMC•  1 ♂; “ Volcán Mombacho. El Progreso.  30-VI-98. JM. Mars”; IEXA. –   Jinotega• 1 ♂; “ El Jaguar Coffee Finca.  XII-3–8– 2005. 4356 ft. D. G. Marqua”; TAMU•  2 ♀♀; “ El Jaguar Coffee Finca.  VI-5–10–2005, el.  4,356 ft.Coll. D. G. Marqua”; VMC•  1 ♂, 2 ♀♀; “ Finca El Jaguar, 32kmNW.  1340m. 13°14’28’’N-86°03’16’’W.  xii-05col D.G. Marqua”; TAMU•  1 ♂; same collection data as for preceding; VMC.   COSTA RICA–  Cartago• 1 ♂; “Turrialba.  650m.  26.Feb.1980. H & A Howden”; TAMU.   Type locality  Nicaragua, Chontales.    Redescription  Major male HEAD. Clypeus bidentate, black anteriorly, bright metallic red, green, or dark metallic blue posteriorly; roughened sculpture. Genae bright metallic red, green, or dark metallic blue; roughened sculpture. Front black, bright metallic red, green, or dark metallic blue on portions adjacent to cephalic horn. Cephalic horn black, curved posteriorly over pronotum ( Figs 2J, 12A–B, D). PRONOTUM. Keel absent in the middle of anterior pronotal margin. Disc triangular, flat, with two distinctly developed tubercles on anterior portion. Triangle bright metallic red, green, or dark metallic blue; becoming black on posterior margin of posterolateral angles; lightly granulate, scabriculous, impunctate. Sides bright metallic red, green, or dark metallic blue; smooth sculpture, scabriculous, with superficially impressed punctures. Lateral lines of pronotal triangle straight. Posterolateral angles widened or slightly acute; projected posteriorly or posterolaterally. Lateral fossae distinctly impressed. Basal fossae obtusely oval, distinctly impressed. Posterior margin sometimes black, with superficially impressed to effaced punctures ( Figs 2J, 12A–B, D). ELYTRA. Striae fine, smooth, scabriculous, not strongly impressed basally; bright red, green, or dark blue; with superficially impressed punctation. Interstriae black, smooth, scabriculous, with almost effaced to effaced punctures. Sutural margin without apical tooth ( Fig. 12A–B, D). PROTIBIAE. Quadridentate with apical spine. TERGITE VIII. Bright metallic red, green, or dark metallic blue; scabriculous; with rough, superficially impressed punctures. Basal margin with setae variable in size. GENITALIA. Right and left lobes of endophallite copulatrix similar in size. Right lobe strongly reduced, obtusely triangular in shape; rounded superiorly. Left lobe obtusely lobed, strongly developed. Central ridge and column similar in size ( Fig. 1N).  Minor male Like the major male, except for the reduction of secondary sexual characters (i.e., cephalic horn, pronotal triangle and tubercles, and pronotal posterolateral angles).  Female Similar to the male, except for the head showing a cephalic trituberculate carina; with conical, nearly aligned tubercles; middle tubercle slightly more developed than lateral tubercles; frons with distinctly impressed punctures; pronotal sculpture smooth, with almost effaced punctures; pronotum almost completely black, becoming posteriorly and laterally bright metallic red, green, or dark metallic blue; pronotal process trituberculate, lacking concavity; pronotal tubercles nearly aligned; with middle tubercle more developed than lateral tubercles; posterior pronotal midline superficially impressed ( Fig. 12C).  Variation Mean length 17.8 mm( 14.7–20.1 mm).  Phanaeus pyroisis the most variable in colour species of the  P. endymionspecies group. The outspoken colour variability of this species was previously outlined by Bates (1886–1889), particularly for the specimens from Nicaragua. Tree typical chromatic morphs were found by us (bright metallic red, Fig. 14A, D; green, Fig. 2J; or dark metallic blue, Fig. 12B), but colour combinations are found and rare specimens has a bright golden sheen.    Distribution  Nicaraguaand north-Caribbean Costa Rica( Fig. 16). The distributions of  P. pyroisand  P. panamensis sp. nov.show an important sympatry area in north Caribbean Costa Rica.    Remarks   Phanaeus pyroisand several closely related species were incorrectly lumped together by previous authors ( Howden & Young 1981; Edmonds 1994; Edmonds & Zídek 2012; Solís & Kohlmann 2012; Chamorro   Fig. 12.  Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH).  et al. 2018, 2019; GBIF Secretariat 2019b). Differences in body colour and the pronotal, elytral and genital morphology were found to confidently diagnose  P. pyroisand all the closely related species. The blue chromatic morph of  P. pyrois( Fig. 12B) was suggested by Edmonds (1994)to be a hybrid with  P. endymion. Nevertheless, blue specimens of  P. pyrois( Fig. 12B) do not share the diagnostic characters with  P. endymion( Figs 1D, 2B, 4). As a consequence, there is no evidence to consider a hybridization between  P. endymionand  P. pyrois. Edmonds & Zídek (2012)suggested that doubtful specimens of “  viridicollis” ( Figs 2J, 12C) were collected in Nicaraguaalong with “normal”  P. pyrois. After revising the doubtful specimens of “  viridicollis” from Nicaragua( Figs 2J, 12C), we disagree with Edmonds & Zídek (2012)and conclude that they incorrectly referred to the green chromatic morph of  P. pyroisas  P. viridicollis. 3093980511 BMNH T. Belt Nicaragua Chontales 41 42 2 2 Chontales lectotype 3093980353 [189,1379,1454,1481] 1998-06-30 TAMU JM. Mars Nicaragua Volcan Mombacho 41 42 1 1 Granada 3093980488 [189,1068,1490,1517] 1998-05-21 Malaise VMC JM. Mars Nicaragua Volcan Mombacho 41 42 1 1 Granada 3093980448 1998-06-30 IEXA JM. Mars Nicaragua Volcan Mombacho 41 42 1 1 Granada 3093980450 2005-12-03 2005-12-08 2005-12-03 TAMU D. G. Marqua Nicaragua El Jaguar Coffee Finca 41 42 1 1 Jinotega 3093980310 2005-06-05 2005-06-10 2005-06-05 VMC D. G. Marqua Nicaragua 1328 El Jaguar Coffee Finca 41 42 2 2 Jinotega 3093980525 2005-12 TAMU D. G. Marqua Nicaragua 1340 13.241112 Finca El Jaguar 21 -86.05445 41 42 3 2 1 Jinotega 3093980517 [525,1106,1634,1660] 2005-12 VMC D. G. Marqua Nicaragua 1340 13.241112 Finca El Jaguar 21 -86.05445 41 42 1 1 Jinotega 3093980530 [189,1251,1700,1727] 1980-02-26 TAMU H & A Howden Costa Rica 650 Cartago 41 42 651 1 Cartago