Geotrupes (Onthotrupes) nuntiatus, Arriaga-Jiménez & Kohlmann & Cruz-García, 2020

Arriaga-Jiménez, Alfonsina, Kohlmann, Bert & Cruz-García, Brenda J., 2020, A predicted new Geotrupes from the mountains of Oaxaca, Mexico, and a description of the male of Geotrupes lobatus (Coleoptera: Geotrupidae), Acta Entomologica Musei Nationalis Pragae 60 (2), pp. 493-508 : 494-499

Geotrupes (Onthotrupes) nuntiatus Kohlmann & Arriaga-Jiménez, sp. nov.

( Figs 1 , 3 , 5 , 10 , 11 )

Type material. HƟΓƟΤΥΡΕ: ♁ (deposited in IEXA; Figs 1a , 2b ), ‘México, Mixteca (Cerro del Águila) Oaxaca / 27-VIII-18 / Coprotrampa / x- - 97.663923° W y- 17.130330° N / Bosque, 3210 m / Arriaga J.A. Col.’ [printed]. AΓΓƟΤΥΡΕ: ♀ ( Fig. 1b ), ‘ México, Mixteca (Cerro del Águila) / Oaxaca / 20-VII-18 / Coprotrampa / x- - 97.685581° W y- 17.114517° N / Bosque, 3012 m / Arriaga J.A. Col.’ [printed]. PΑ*©ΑΤΥΡΕඌ (768 specimens): MEXICO: OΑΧΑർΑ: 17-VI-18 Mixteca (Cerro del águila) Oaxaca C.D. perro. 17.119718° -97.684495° 3065m Bosque AAJ col (2 ♁♁ CZUG, 2 ♀♀ CMN, 2 ♁♁ NMPC). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca C.D. perro 17°7’35.5” -97°39’55.65” 3135m Matorral AAJ col (10 ♁♁ IEXA, MNHN, 2 ♀♀ IEXA, NHMUK). 21-VII- 18 Mixteca (Cerro del águila) Oaxaca C.D. vaca 17.12301° -97.67303° 2967m Bosque AAJ col (1 ♁). 19-VII-18 Mixteca (Cerro del águila) Oaxaca C.D. 17.126010° -97.681780° 2820m Matorral AAJ col (5 ♁♁ CMN, 4 ♀♀ MNHN, NHMUK). 17-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.119661° -97.684522° 3066m Bosque AAJ col (3 ♁♁ CZUG, IEXA). 13-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130087° -97.708348° 2861 m Bosque AAJ col (1 ♀). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.13123° -97.70777° 2807 m Bosque AAJ col (1 ♁ UAEH, 2 ♀♀ CZUG).27-VIII- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.13232° -97.70776° 2791 m Bosque AAJ col (3 ♀♀ IEXA). 01-IX-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.183022° -97.727770° 2234m Bosque AAJ col (1 ♁ MNHN, 1 ♀ NMPC). 30-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.152799° -97.713897° 2642m Bosque AAJ col (1 ♁ CMN). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.132831° -97.707548° 2839m Bosque AAJ col (1 ♁ CMN). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.133220° -97.707226° 2844 m Bosque AAJ col (1 ♁ FEST). 27-VIII- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.133220° -97.707226° 2844 m Bosque AAJ col (4 ♁♁ CMN, 4 ♀♀ CZUG, IEXA, 2 ♀♀ NMPC).20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.133487° -97.70781° 2804m Bosque AAJ col (1 ♁). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.133487° -97.706736° 2858m Bosque AAJ col (1 ♁, 1 ♀ IEXA). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.133381° -97.706166° 2849m Bosque AAJ col (4 ♁♁ CMN, 1 ♀ CMN). 22-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.123113° -97.673899° 3136 m Bosque AAJ col (1 ♁ MNHN, 1 ♀ NMHUK). 20-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.152902° -97.715444° 2624 m Bosque AAJ col (4 ♁♁ MNHN, NHMUK, 2 ♁♁ NMPC). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.105464° -97.705154° 2810 m Bosque AAJ col (1 ♀ FEST). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.110796° -97.687456° 3010 m Bosque AAJ col (1 ♁, 2 ♀♀ CMN). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111180° -97.687293° 3004 m Bosque AAJ col (1 ♀ FEST). 29-VIII- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111180° -97.687293° 3004 m Bosque AAJ col (1 ♁ CZUG). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111627° -97.686797° 3015m Bosque AAJ col (1 ♁ CMN). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111627° -97.686797° 3015m Bosque AAJ col (1 ♁ CMN, 1 ♀ CMN). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.113481° -97.685020° 3002m Bosque AAJ col (2 ♁♁ UAEH, 1 ♀ CZUG). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.113481° -97.685020° 3002m Bosque AAJ col (1 ♁ FEST). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.113481° -97.685002° 3002 m Bosque AAJ col (2 ♁♁ UAEH). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114517° -97.685581° 3012m Bosque AAJ col (1 ♁ FEST, 2 ♀♀ JBLPM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114517° -97.685581° 3012m Bosque AAJ col (4 ♁♁ CZUG, IEXA, 7 ♀♀ CZUG, JBLPM, IEXA). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114517° -97.685581° 3012m Bosque AAJ col (2 ♁♁ CMN). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114061° -97.685462° 3023 m Bosque AAJ col (2 ♀♀ FEST). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125511° -97.681109° 3158m Bosque AAJ col (4 ♁♁ FEST, IEXA, 2 ♀♀ FEST, JBLPM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125511° -97.681109° 3158m Bosque AAJ col (1 ♀ UAEH). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125511° -97.681109° 3158m Bosque AAJ col (3 ♁♁ UAEH, 6 ♀♀ CZUG, UAEH). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125921° -97.681559° 3168 m Bosque AAJ col (7 ♁♁ IEXA, JBLPM, NHMUK, UNAM, 5 ♀♀ CMN, FEST, JBLPM). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125921° -97.681559° 3168 m Bosque AAJ col (5 ♁♁ JBLPM, 1 ♀ FEST, JBLPM). 23-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125921° -97.681559° 3168 m Bosque AAJ col (9 ♁♁ CZUG, FEST, UAEH, 7 ♀♀ FEST, UAEH, 1♀ NMPC). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125921° -97.681559° 3168 m Bosque AAJ col (4 ♁♁ UNAM, 5 ♀♀ FEST, JBLPM, UNAM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130874° -97.664017° 3234 m Bosque AAJ col (1 ♁ CZUG, 3 ♀♀ UAEH). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130874° -97.664017° 3234 m Bosque AAJ col (2 ♁♁ FEST, 2 ♀♀ CMN, FEST). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130874° -97.664017° 3234 m Bosque AAJ col (1 ♁, 8 ♀♀). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130874° -97.664017° 3234 m Bosque AAJ col (3 ♁♁ CMN, 6 ♀♀ CMN, MNHN, UNAM). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130330° -97.663923° 3219 m Bosque AAJ col (3 ♁♁ FEST, 1 ♀ FEST). 22-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130330° -97.663923° 3219 m Bosque AAJ col (8 ♁♁ CZUG, UAEH, 7 ♀♀ CZUG). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130330° -97.663923° 3219 m Bosque AAJ col (13 ♁♁ CMN, IEXA, UNAM, 8 ♀♀ IEXA, UAEH, UNAM, 3 ♀♀ NMPC). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130330° -97.663923° 3219 m Bosque AAJ col (7 ♁♁ CZUG, IEXA, 8 ♀♀ CZUG, IEXA, UNAM, 1 ♁ NMPC). 20-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129932° -97.663673° 3222m Bosque AAJ col (1 ♀). 13-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129932° -97.663673° 3222m Bosque AAJ col (1 ♁ UAEH, 3 ♀♀ UAEH). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129932° -97.663673° 3222m Bosque AAJ col (6 ♁♁ CMN, 2 ♀♀ CMN). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.663794° 3220m Bosque AAJ col (1 ♀). 19- VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.663794° 3220m Bosque AAJ col (1 ♀). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.663794° 3220m Bosque AAJ col (1 ♁, 8 ♀♀). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.663794° 3220m Bosque AAJ col (5 ♁♁, 6 ♀♀). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.663794° 3220m Bosque AAJ col (9 ♁♁, 11 ♀♀, 1 ♁ NMPC). 20-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128973° -97.663948° 3217m Bosque AAJ col (9 ♁♁, 13 ♀♀). 27- VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128973° -97.663948° 3217m Bosque AAJ col (10 ♁♁, 13 ♀♀). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128973° -97.663948° 3217m Bosque AAJ col (10 ♁♁, 12 ♀♀). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (1 ♀ CMN). 31-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (6 ♁♁ CMN, IEXA, UNAM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (2 ♁♁, 10 ♀♀). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (4 ♁♁ CMN, IEXA, UNAM, 5 ♀♀ CMN). 27-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (1 ♁). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128176° -97.661997° 3232m Matorral AAJ col (16 ♁♁, IEXA, UNAM, 6 ♀♀). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.127822° -97.662464° 3229 m Matorral AAJ col (8 ♁♁, 10 ♀♀ CMN, IEXA). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.127822° -97.662464° 3229 m Matorral AAJ col (2 ♁♁ CMN, 5 ♀♀ CMN, UNAM). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.127822° -97.662464° 3229 m MatorralAAJ col (14 ♁♁ CEMT,FEST, IEXA, UNAM, 11 ♀♀ CMN, FEST, JBLPM). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.127822° -97.662464° 3229 m Matorral AAJ col (2 ♁♁ MNHN, 4 ♀♀ MNHN, 2 ♀♀ NMPC). 29- VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.127822° -97.662464° 3229 m Matorral AAJ col (2 ♁♁, 1♀ NMPC) 20-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128058° -97.662982° 3241 m Bosque AAJ col (6 ♁♁ MNHN, NHMUK, 4 ♀♀ MNHN, NHMUK). 25-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128058° -97.662982° 3241 m Bosque AAJ col (7 ♁♁ CMN, MNHN, 18 ♀♀ CMN, MNHN, UNAM). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128058° -97.662982° 3241 m Bosque AAJ col (2 ♁♁). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128058° -97.662982° 3241 m Bosque AAJ col (2 ♁♁ CMN, 3 ♀♀ CMN). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128267° -97.663421° 3242m Bosque AAJ col (6 ♁♁ MNHN, NHMUK, 7 ♀♀ NHMUK, UNAM). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128267° -97.663421° 3242m Bosque AAJ col (5 ♁♁ CMN, 11 ♀♀ CMN, IEXA). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128860° -97.663403° 3236 m Bosque AAJ col (2 ♁♁ MNHN, UAEH, 2 ♀♀ NHMUK, 2 ♁♁ NMPC). 21-VI- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128860° -97.663403° 3236 m Bosque AAJ col (3 ♁♁ FEST). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128860° -97.663403° 3236 m Bosque AAJ col (4 ♁♁ UAEH, 13 ♀♀ CZUG, IEXA, 1 ♀ NMPC). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129932° -97.663673° 3222m Bosque AAJ col (3 ♁♁ UAEH, 3 ♀♀ CZUG). 19- VI-18 Mixteca (Cerro del águila) Oaxaca C.D. 17.13232° -097.70776° 2791 m Bosque AAJ col (12 ♁♁ CMN, MNHN,UNAM, 11 ♀♀ JBLPM, MNHN, NHMUK, UNAM, 1 ♁ NMPC). 1-IX-18 Mixteca (Cerro del águila) Oaxaca C.D. 17.13232° -097.70776° 2791 m Bosque AAJ col (3 ♁♁ CZUG, 1 ♀). 28-VI-17 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17°05’33.9” -97°39’18.1” 2744m Bosque de pino-encino AMT col (1♀ UNAM, 1 ♁ NMPC). 13-VII-17 Mixteca (Cerro del águila) Oaxaca Micetotrampa 17°03’57.1” -97°38’13.5” 2629m Bosque de pino-encino AMT col (1♀ UNAM). 13-VII-17 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17°05’16.9” -97°39’42.2” 2535m Bosque de pino -encino AMT col (2 ♀♀ UNAM, 1 ♁ NMPC). 28-IX-17 Mixteca (Cerro del águila) Oaxaca Micetotrampa 17°05’18.5” -97°39’47.0” 2543m Bosque de pino-encino AMT col (2 ♁♁ UNAM, 1♀ UNAM). 13-VII-17 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17°05’18.5” -97°39’47.0” 2543m Bosque de pino-encino AMT col (2 ♀♀ JBLPM). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125511° -97.681109° 3158 Bosque AAJ col (2 ♁♁). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128267° -97.663421° 3242m Bosque AAJ col (11 ♁♁, 12 ♀♀). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128973° -97.663948° 3217m Bosque AAJ col (8 ♁♁, 3 ♀♀). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128267° -97.663421° 3242m Bosque AAJ col (2 ♁♁, 1 ♀). 23-VIII- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.128267° -97.663421° 3242m Bosque AAJ col (3 ♁♁, 4 ♀♀). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125921° -97.681559° 3168m Bosque AAJ col (2 ♁♁ CEMT, NHMUK, 2 ♀♀ CEMT). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114517° -97.685581° 3012m Bosque AAJ col (3 ♁♁ NHMUK, 4 ♀♀ CEMT). 23-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.132831° -97.707548° 2839m Bosque AAJ col (4 ♁♁ NHMUK, 3 ♀♀ CEMT). 23-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130874° -97.664017° 3234 m Bosque AAJ col (4 ♁♁ NHMUK, 3 ♀♀ CEMT). 23-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.125511° -97.681109° 3158 m Bosque AAJ col (3 ♁♁ NHMUK, 1 ♀ CEMT). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111627° -97.686797° 3015 m Bosque AAJ col (2 ♁♁ NHMUK, 3 ♀♀ CEMT). 21-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114517° -97.685581° 3012m Bosque AAJ col (1 ♀ NHMUK). 24-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.152430° -97.714620° 2661m Bosque AAJ col (1 ♁ NHMUK). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111180° -97.687293° 3004 m Bosque AAJ col (2 ♁♁ CEMT, 2 ♀♀ CEMT). 20-VI-18 Mixteca (Cerro del águila) Oaxaca C.D. exc.perro 17.159213° -97.721293° 2520m Bosque AAJ col (4 ♁♁ CEMT, 2 ♀♀ UNAM). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.110796° -97.687456° 3010 m Bosque AAJ col (3 ♁♁ CEMT, 5 ♀♀ UNAM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.130330° -97.663923° 3219 m Bosque AAJ col (1 ♁ CEMT). 23-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.112194° -97.685904° 3024 m Bosque AAJ col (1 ♁ CEMT, 1 ♀ CEMT). 01-IX- 18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.152430° -97.714620° 2661m Bosque AAJ col (1 ♁ CEMT). 27-VI-17 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17°06’11.1” -97°39’23.2” 2732m Bosque de pino-encino AMT col (4 ♁♁ CEMT, IEXA, UNAM, 1♀ UNAM). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111180° -97.687293° 3004 m Bosque AAJ col (1 ♀ CEMT). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111627° -97.686797° 3015 m Bosque AAJ col (2 ♀♀ CEMT). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114061° -97.685462° 3023 m Bosque AAJ col (1 ♀ CEMT, 2 ♁♁ 1 ♀ NMPC). 27-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.114061° -97.685462° 3023 m Bosque AAJ col (1 ♀ NMPC). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.111627° -97.686797° 3015 m Bosque AAJ col (1 ♀). 20-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.13072° -97.70813° 2825 m Bosque AAJ col (2 ♀♀ CEMT). 19-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.113481° -97.685020° 3002m Bosque AAJ col (1 ♀ UNAM). 24-VI-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.151172° -97.714956° 2633 m Bosque AAJ col (1 ♀ UNAM). 24-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.153153° -97.713830° 2632 m Bosque AAJ col (1 ♀ UNAM). 17-VI-18 Mixteca (Cerro del águila) Oaxaca C.D. exc. vaca 17.123698° -97.674714° 3154 m Bosque AAJ col (1 ♀ UNAM). 29-VIII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.13011° -097.10835° 2836 m Bosque AAJ col (1 ♀ UNAM, 2 ♀♀ NMPC). 23-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.129433° -97.662464° 3220m Bosque AAJ col (3 ♁♁, 8 ♀♀). 13-VII-18 Mixteca (Cerro del águila) Oaxaca Coprotrampa 17.13011° -097.10835° 2836 m Bosque AAJ col (1 ♁ UAEH, 5 ♀♀ CMN, UAEH). 27-VI-18 Mixteca (Cerro del águila) Oaxaca C.D. perro. 17.119718° -97.684495° 3065m Bosque AAJ col (1 ♁ MNHN). 20-VI-18 Mixteca (Cerro del águila) Oaxaca C.D. perro 17.152568° -97.713997° 2629m Bosque AAJ col (1 ♁ MNHN). 22-VII- 18 Mixteca (Cerro del águila) Oaxaca C.D. exc. Borrego 17.123113° -97.673899° 3136 m Bosque AAJ col (1 ♀). (Localities with no indication represent specimens deposited at IEXA).

Body measurements were made to the nearest 0.1 mm using an ocular micrometer with a Zeiss Stemi DV4 stereoscope, Jena, Germany. Photographs were taken by: Susana Guzmán-Gómez (Instituto de Biología, Universidad Nacional Autónoma de México, IBUNAM, Mexico City, Mexico) using a Zeiss AXIO Zoom V16 microscope, a Zeiss AxioCam MRc5 camera, and the Zeiss efficient navigation multifocal technology program (Zeiss, Jena, Germany); BJCG using a multifocus digital microscope Leica Z16 APOA; Alfonso Aceves (INECOL) using a Canon T 2i camera, extension tubes, a 100 mm macro, and an external Canon flash; and photographs of material from the CMNC were taken by AAJ using a Leica Z16 system and LAS software for image stacking. The distribution maps were made using QGIS desktop 2.14.19.

Diagnosis. Dorsally shining ( Figs 1 a–b); antennal club yellowish ( Figs 1 a–b); mandibles lacking a very pronounced rounded lobe ( Figs 1 a–b); legs black; male protibiae short and straight with apical teeth flattened ( Fig. 3a ); male profemora without a transverse ridge at its base and not having profemora ventrally excavated near coxae; pronotum, scutellum and elytral intervals shining, green, golden or blue with coppery reflections ( Fig. 1a ); posterior pronotal margin distinct in front of scutellum ( Fig. 1a ); elytral intervals convex ( Fig. 1a ); male procoxa forming a slender saddle-like structure at point of junction with trochanter; male mesoventrite with a distinct, projected keel in front of mesocoxae; aedeagus like Figs 1 c–f.

Description of holotype. Male ( Figs 1a , 3a ), length 18.4 mm, greatest width 9.4 mm. Color dorsally golden-green with strong reddish-coppery pronotal reflections. Antennal club reddish-brown, unmodified; external face of terminal antennomere with a faint U-shaped furrow. Clypeus abruptly arcuate anteriorly; disc rugose and with small, conical tubercle posteriorly. Edge of ocular canthus obtusely angulate anteriorly. Pronotum with small, non-punctate depression by midline just behind anterior margin; posterior margin vaguely indicated in front of scutellum, otherwise absent; disc coarsely punctate anteriorly and laterally. Scutellum impunctate, midline impressed, sides arcuate on anterior third. Elytra distinctly striate, first stria terminated by scutellum, extending basally no more than one-third length of scutellum; striae shallowly, finely punctate, crenate. Protibia with apical tooth bent inwardly ( Fig. 3a ), apex shallowly emarginate; additional five teeth on outer margin unmodified; flattened ventral or inner surface with two distinct conical teeth, apical tooth slightly smaller; a third-minute tooth often evident on basal side of larger proximal tooth; mesotibia with three complete, transverse carinae on outer surface; metatibia with four complete, transverse carinae on outer surface; metafemur with tooth near trochanter lacking. Mesoventrite with small, distinct keel in front of mesocoxae. Aedeagus as in Figs 1 c–f.

Description of allotype. Female ( Fig. 1b ), length 19.1 mm, greatest width 9.7 mm. Similar to male except in the following respects: protibia with seven unmodified teeth on outer edge, apical tooth simple ( Fig. 1b ); ventral surface of protibia lacking two large conical teeth; apical spur longer, thicker and bent inward ( Fig. 1b ); scutellum border strigose; coxa not forming a ridge at juncture with trochanter.

Variation. Body length 11.6–19.2 mm. Maximum width 6.9–10.1 mm. Coloration ranges from bright metallic light green to a bright metallic light blue, to a bright metallic green-gold to gold with reddish-coppery reflections. Middle of thorax can go from almost a smooth surface to a faint middle-line, to a faint middle-line with punctures. Differential diagnosis. For the differential diagnosis, see the key below.

Etymology. From the Latin word nuntiatus (part. masc. nom. sing.), meaning the announced one. The name derives from the fact that a species distribution model for the Last Glacial Maximum in Oaxaca predicted this species existence (A*©*©ΙΑǤΑ-JΙΜḖΝΕΖ et al. 2020).

Collection circumstances. The species has been collected in traps baited with human dung and mushrooms (Agaricales and Boletales), as well as directly under sheep, cow, and dog dung. The collections were made from June to September in the “Cerro del Águila” ( Fig. 5 ) in the Sierra de Tlaxiaco and communities around it in the Mixteca shield of Oaxaca, Mexico. It is part of the Sierra Madre del Sur, with a mountain range from 2.200 to 3.250 m a.s.l. This area is characterized by pine, oak and pine-oak forests, with some areas with shrublands near the communities. This mountain has a sub-humid climate with heavy rains in the summer, with mean annual temperatures ranging from 10° to 16°C and medium precipitations oscillating from 800 to 1.500 mm per year (IΝΕǤΙ 2008).

Taxonomic notes. The new species, G. nuntiatus sp. nov., is veritably halfway between G. pecki and G. viridiobscurus . The aedeagus ( Figs 1 c–f) is more similar to G. pecki ( Figs 8 c–f) than to G. viridiobscurus ( Figs 9 c–f). However, the beetle’s external appearance ( Figs 1 a–b) is indistinguishable from G. viridiobscurus ( Figs 9 a–b), except for the coloration, where the specimens of G. nuntiatus sp. nov. are very bright, ranging from green, to blue or golden-black with a reddish-coppery tint versus the specimens of G. viridiobscurus being a dull bluish or greenish-black.

Color as a character has been much maligned in taxonomy. However, HΙΓΓ (2017) has now proposed that the mitonuclear species compatibility concept is the best current method for diagnosing species. This mitonuclear compatibility leads, among other things, to the evolution of species-typical coloration as part of speciation reinforcement processes. Thus, species-typical coloration evolves as a signal of coadapted mitochondrial-genotype genes. HΙΓΓ (2017) presents as a case study the nearly perfect agreement between bird species boundaries according to mtDNA types and those drawn according to species-typical songs or coloration. Therefore, coloration might be a valid taxonomic character, which should be used judiciously and concomitantly with other concordant characters.

Figures

Fig. 1. Geotrupes nuntiatus Kohlmann & Arriaga-Jiménez, sp. nov.: a – male dorsal view (17.13087°N, -97.66402°W); b – female dorsal view (17.129932°N, -97.663673°W); c–f – male genitalia: c – in dorsal view, d – in ventral view; e–f – lateral views. Scale bars = 5 mm (a–b), 2 mm (c–f).

Fig. 3. Morphological detail of male protibiae: a – Geotrupes nuntiatus Kohlmann & Arriaga-Jiménez, sp. nov.; b – G. lobatus Howden, 1974; arrow pointing at the apical tooth on the profemur. Scale bar = 2 mm.

Fig. 5. Map showing the known distribution of Geotrupes lobatus Howden, 1974, G.sallei Jekel, 1866, and G.nebularum Howden, 1974 (upper map) and the known distribution of G. nuntiatus Kohlmann & Arriaga-Jiménez, sp. nov., G. viridiobscurus Jekel, 1866, and G. pecki Howden, 1974 (lower map).

Fig. 10. Projection of potential distribution for Geotrupes viridiobscurus Jekel, 1866 and G. pecki Howden, 1974 in the state of Oaxaca, representing the estimated climatic range at the Hueyatlaco-I glaciation scenario (20–17.5 kyr ago), based on the actual distribution of both species. Areas in green (1.265–2.390 m a.s.l.) correspond to the modern temperature range of G. viridiobscurus (orange locality points, Sierra Norte) and G. pecki (yellow locality points, Sierra Sur). Areas in blue are too cold and areas in yellow are too warm. Areas in blue are very similar to the present-day projected climatic distribution (2.300–3.400 m a.s.l.). The green points represent the localities of G. nuntiatus Kohlmann & Arriaga-Jiménez, sp. nov., the new species being described in this study and belonging to this species group. It falls within the modelled distribution area (modified from A*©*©ΙΑǤΑ-JΙΜḖΝΕΖ et al. 2020).

Fig. 11. Native vegetation (pine forest and montane shrubland) at the top of Cerro del Águila, Mixteca, Oaxaca.

Fig. 2. Geotrupes lobatus Howden, 1974: a – male dorsal view (16.16096°N, -96.24452°W); b – female dorsal view (16.16096°N, -96.24452°W); c–f – male genitalia: c – in dorsal view, d – in ventral view; e–f – lateral views. Scale bars = 5 mm (a–b), 2 mm (c–f).

Fig. 8. Geotrupes pecki Howden, 1974: a – male dorsal view (16.17325°N, -96.24708°W); b – female dorsal view (16.12138°N, -96.45222°W); c–f – male genitalia: c – in dorsal view, d – in ventral view; e–f – lateral views. Scale bars = 5 mm (a–b), 2 mm (c–f).

Fig. 9. Geotrupes viridiobscurus Jekel, 1866: a – male dorsal view (17.44361°N, -96.50361°W); b – female dorsal view (16.160961°N, -96.244523°W); c–f – male genitalia: c – in dorsal view, d – in ventral view; e–f – lateral views. Scale bars = 5 mm (a–b), 2 mm (c–f).