Griburius montezuma ( Suffrian, 1852 )

Sassi, Davide, 2023, Revision of the Griburius scutellaris (Fabricius, 1801) species group (Coleoptera: Chrysomelidae: Cryptocephalinae), Zootaxa 5315 (6), pp. 501-548 : 532-536

Griburius montezuma ( Suffrian, 1852)

( Figs 9 ; 13c–d )

Scolochrus montezuma Suffrian, 1852: 116 (original description); Suffrian, 1858: 389 (taxonomic notes); Jacoby 1880: 59 (taxonomic notes).

Griburius montezuma: Crotch, 1873a: 32 (taxonomic notes); Crotch, 1873b: 96 (catalogue); Horn, 1895: 229 (catalogue); Clavareau, 1913: 90 (catalogue); Beamer, 1926: 209 (biological notes); Blackwelder, 1946: 640 (catalogue); Pallister, 1953: 24 (taxonomic notes); Riley et al.: 2001: 12 (taxonomic notes); Riley et al., 2003: 154 (catalogue); Clark et al., 2004: 139 (biological notes); Andrews & Gilbert, 2005: 96 (catalogue); Chaboo et al., 2016: 157 (biological notes); Niño-Maldonado et al., 2016: 104 (catalogue); Sánchez-Reyes et al., 2015: 528 (catalogue); Sánchez-Reyes et al., 2016: 54 (catalogue); ( Ordóñez-Reséndiz & López-Pérez, 2021: 90 (catalogue).

Griburius speciosus LeConte 1867 (original description): 56; Crotch, 1873a: 32 (as syn. of G. montezuma , taxonomic notes); Crotch, 1873b: 96 (as syn. of G. montezuma , catalogue); Clavareau, 1913: 90 (as syn. of G. montezuma , catalogue); Blackwelder, 1946: 640 (as syn. of G. montezuma , catalogue); ( Ordóñez-Reséndiz & López-Pérez, 2021: 90 (as syn. of G. montezuma , catalogue).

Scolochrus speciosus: Jacoby 1880: 59 (as syn. of G. montezuma , taxonomic notes).

Scolochrus suffriani Jacoby 1880: 60 (original description); Jacoby 1889: 126 (taxonomic notes); Schaeffer 1906: 233 (as syn. (“chromatic form”) of G. montezuma , taxonomic notes).

Griburius montezuma suffriani: Clavareau, 1913: 90 (catalogue); Blackwelder, 1946: 640 (catalogue); Pallister, 1953: 24 (taxonomic notes). Syn. restored.

Schaeffer (1906) proposed the synonymy of G.suffriani , justifying his judgment through an in-depth discussion of the intraspecific variability of G. montezuma . Despite this, in subsequent catalogues ( Clavareau, 1913; Blackwelder, 1946) the name suffriani is given an ambiguous rank of “var.”, which should be considered as subspecific ( ICZN, 1999, Art. 45.6.3). Additionally, Pallister (1953) expressly attributed the rank of subspecies to G. suffriani . Only in Riley et al. (2003), as far as I could ascertain, does G. suffriani reappear, once again without comments, listed as a junior synonym of G. montezuma . Fully confirming Schaeffer’s judgment on the great chromatic variability of G. montezuma , given the current ambiguity of the nomenclatural framework, and given the fact that the last critical discussion on the validity of the name was the one of Pallister (1953) I consider it appropriate to formally reaffirm, the synonymy of the name G. suffriani with respect to G. montezuma , to clarify the status of this name in the context of a taxonomic work.

Scolochrus cazicus Suffrian, 1852: 119 (original description); Suffrian, 1858: 389 (taxonomic notes); Suffrian, 1866: 201 (taxonomic notes); Jacoby 1880: 60 (taxonomic notes).

Griburius cazicus Clavareau, 1913: 89 (catalogue); Blackwelder, 1946: 639 (catalogue); Ordóñez-Reséndiz & López-Pérez, 2021: 90 (catalogue). Syn. nov.

Types. Griburius montezuma: Suffrian (1852) stated that he had a single female available for the study (“from Berlin Museum”). In MNHUB a female is housed, which matches the description, therefore it is here accepted as the holotype of the species. HOLOTYPE: ♀, pinned, // “23992” [white label, printed] // “Montezuma Suffr. Mexico Koppe” [blue label, handwritten] // “ Griburius montezuma ( Suffrian, 1852) ( Scolochrus montezuma ) HOLOTYPUS D. Sassi det.” [red label, printed] // (MNHUB).

G. speciosus : this taxon was described by John Lawrence LeConte (1867) on two specimens from “Raton Mountain”. A single specimen is housed in MCZ, which matches the description and is here designated as lectotype: ♀, glued, // “ G. montezumae Suff. speciosus Lec. ” [white label, handwritten] // “N. M.” [white label, handwritten] // Type 4288.” [red label, partly handwritten] // “ Griburius speciosus LeConte, 1867 LECTOTYPUS D. Sassi det.” [red label, printed] // Griburius montezuma Suffrian, 1852 D. Sassi det, 2022 [white label, printed] // (MCZ).

G. suffriani: Jacoby (1880) in the original description indicates that a single male was available for the study, therefore a specimen in BMNH which perfectly meets the original description is here accepted as the holotype of the species. HOLOTYPE: ♁, pinned, // “Tehuantepec” [white label, printed] // “ Mexico Salle Coll,” [white label, printed] // “298” [white label, printed] // “ Scolochrus suffriani Jacoby ” [blue label, handwritten] // “Type Sp. figured” [white label, printed] // “B.C.A., Col. VI,1. Scolochrus Suffriani Jac. ” [white label, partly handwritten] // “Type H. T.” [white and red label, printed] // “ Griburius suffriani ( Jacoby, 1880) ( Scolochrus suffriani ) HOLOTYPUS D. Sassi det.” [red label, printed] // Griburius montezuma Suffrian, 1852 D. Sassi det, 2022 [white label, printed] // (BMNH).

G. cazicus : A female in BMNH matches the original description and, therefore, it is here designated as lectotype of the taxon since Suffrian did not indicate the number of specimens available to him. LECTOTYPE: ♀, pinned, // “ Scolochrus cazicus Suff [unreadable word] Chevr Yucatan ” [white label, handwritten] // “E. Coll. Chev. t ” [white label, printed] // “64” [white label, printed] // “67.56” [white label, printed] // “ Griburius cazicus Suffrian, 1852 ( Scolochrus cazicus ) LECTOTYPUS D. Sassi det.” [red label, printed] // Griburius montezuma Suffrian, 1852 D. Sassi det, 2022 [white label, printed] // (BMNH).

Type localities. G. montezuma : Mexico. G. speciosus: Raton Mountain (New Mexico, USA). G. suffriani: Tehuantepec ( Mexico) . G. cazicus : Yucatan ( Mexico)

Additional material examined. EL SALVADOR: Cabañas: Tejutepeque 19.VI.2021 (1, GBIF) . GUATEMALA: Baja Verapaz: San Jeronimo 6.VII.2020 (1, GBIF) . MEXICO: BAJA CALIFORNIA: 6 km Hwy 1@ km 54 rd Rumal de los Naranjos 26.IX.2003 (1, BYU) ; Ramal de los Naranjas 0–6 km W Hwy 1 26.IX.2003 (1, BYU) ; Basse Californie Diguet 178–95 (2, MNHN) . BAJA CALIFORNIA SUR: Cabo San Luca 21.VI.1979 (1, TAMU) ; 9 mi. N Cabo San Lucas Blacklight 15.IX.1988 (2, ERPC) ; S Todos Santos Rancho El Cachania 30.IX.1989 (1, ERPC) ; 6, 5 mi. S El Pescadero (1, FSCA) . CHIAPAS: Parque Nacional El Guacero 23.V.1987 (1, ERPC) . CHIHUAHUA: Majalca Rd. 6000’ 35 mi. NW Chihuahua 14.VI.1961 (1, CNCI) . DURANGO: La Registrillo 15 Km SE Durango 7.VI.1995 (1, USNMNH) ; 25 mi. W Durango 3.VII.1964 (1, CNCI) ; 5 mi. E Durango 6500’ 20.VI.1964 (1, CNCI) ; 5 mi. W Durango 16.VII.1964 (1, CNCI) . HIDALGO: Tezontepec de Aldama 22.V.2022 (1, GBIF) . GUERRERO: 6 mi WE Taxco 5600’ 14.VIII.1974 (1, ERPC) ; 38 km W Iguala 16.VI.1995 (1, USNMNH) ; 3 km NW Cacahuamilpa 1500m 2.VII.1992 (1, NHMB) ; 2.1 mi NW of Cacahuamilpa 27.VI.1983 (1, TAMU) ; 3.3 km Cacahuamilpa 1495m 2.VII.1992 (2, ERPC) ; 20 mi N Iguala 22.VIII.1958 (1, CNCI) ; 6 mi E Xochipala 13.VII.985 & 18.VII.1985 (9, TAMU) ; 6 mi E of Xochipala 13.VI.1985 (2, TAMU) ; Agua de Correa, Zihuatanejo de Azueta 30.V.2021 (1, GBIF) ; La Villa Ayutla de los Libres 27.VII.2019 (1, GBIF) ; Los Troncones, Troncones 1.VIII.2022 (1, GBIF) . JALISCO: Chamela Estcn UNAM 1.X.1991 (1, TAMU) ; Chamela Vic. ESTC UANM 9.VII.1993 (2, TAMU) ; Chamela Biol. Exp. Sta. UNAM 200’ 9.VIII.1982 (1, ERPC) ; Vulkan Colima Esperanza 1000m 1918 (1, ZSM) . MORELOS: 4.4 mi. E Cuernavaca malaise trap 27.VII.1976 (1, TAMU) ; Cuernavaca 1.VIII.1944 (1, USNMNH) ; Cacahuamilpa 1495m 2.VII.1992 (6, NHMB) ; Xochicalco Ruins 14.VII.1995 (1, USNMNH) ; Cuatla 13.VII.1946 (1, AMNH) ; Laguna de Tequesquitengo 950m 1946 (1, MNHN) ; Xochitepec 13.VIII.2018 (1, GBIF) . NUEVO LEON: Santa Rosa Canyon 28 km W Linares 31.V.1987 (1, FSCA) . NAYARIT: Rosamorada 27.VII.2020 (1, GBIF) . OAXACA: 6 mi W of Jalapa del Marques at light 23.VI.1973 (3, TAMU) ; Tehuantepec (4, USNMNH) ; 10 mi N Salina Cruz nr shore 16.I.1982 (1, ERPC) ; 6 mi NE Mitla 20.VII.1985 (2, TAMU) ; Puerto Angel 20.VII.1963 (1, USNMNH) ; 5 mi NW of Huajuapan 28.VII.1981 (1, TAMU) ; Juchitán de Zaragoza 13.VIII.2020 (1, GBIF) . PUEBLA: 11.8 Mi. NW Izucar de Matamoros 13.VI.1974 (1, TAMU) ; 10 km NW Petlalcingo 5.VII.1992 (1, NHMB) ; 6 mi. NW Acatlan 22.VII.1974 (1, FSCA) ; 1.1 mi W Acatlan at light 13.VI.1974 (1, TAMU) ; 14.6 mi. N Tchuitzingo 9.VIII.1980 (1, TAMU) . QUERÉTARO: Fraccionamiento Vista Real, Corregidora 15.VI.2021 (1, GBIF) ; Cadereyta de Montes , 24.VII.2017 (1, GBIF) . QUINTANA ROO: Luis Donaldo Colosio, Playa del Carmen 18.VII.2019 (1, GBIF) ; Solidaridad 6.VII.2022 (1, GBIF) . SAN LUIS POTOSÍ: San Luis Potosí , 2.VII.2022 (1, GBIF) ; 73 km N San Luis Potosí 2.VI.1987 (1, TAMU) ; 21 km N Entronque el Hulzache 2.VI.1987 (1, FSCA) . SINALOA: Mazatlán 5.IX.2020 (1, GBIF) . SONORA: N Hermosillo 25.V.1961 at light (1, CNCI) ; 17 km S Moctezuma 30.VIII.1980 at light (2, TAMU) ; Micro Cerro Prieto 15 km E Navajoa 400m 24.VII.2008 (1, BYU) ; 5 mi. W Guaymas 21.IV.1949 (1, AMNH) . VERACRUZ: Cotaxtla Exp. Sta. 8.VIII.1962 Acacia cornigera (5, USNMNH) . YUCATÁN: Motul 11.VII.1952 (2, AMNH) . U.S.A.: ARIZONA: Cochise Co.: 3 mi NW Portal 11.VII.1972 (1, TAMU) ; Copper Canyon Huachuca Mts. 2.VIII.1991 (2, BYU) ; Dragon Mountains Middlemarch Pass 29.VI.1997 (1, BYU) ; Cochise Stronghold 7.VII.1986 (1, BYU) ; Chiricahua Mts. 24.VII.1955 & 17.VII.1957 & 23, VII.1959 (3, BYU) ; Chiricahua Mts. 23.VI.1997 (1, ERPC) . Coconino Co.: Oak Creek 10 Km N Sedona 16.VII.2004 (1, NMV) . Gila Co.: Pinal Mts. Ocehouse Can. 26.VI.1979 1414m at light (1, TAMU) ; Graham Co.: Aravaipa Can. 21.VI.1976 at light (1, TAMU) ; Galiuro Mts. High Cr. 20.VII.1978 at light (1, TAMU) . Maricopa Co.: Hassayampa River Preserve 24.VI.2022 (1, GBIF) . Mohave Co.: 1 mi S Haekberry Night Light 3.VIII.1970 (1, BYU) ; 2 mi. E Hualapai Mountain Road on Rt. 259 15.VII.2011 (1, BYU) . Navajo Co.: 3 miles N Snowflake 13.VI.1967 (1, TAMU) . Pima Co.: Catalina Mts. Sabino Cyn. 1.VI. 1980 (2, FSCA) ; 4 mi S Arivaca Fraguita Wash 10.VII.1977 at light (1, TAMU) ; Tucson 13.VI.1982 (1 TAMU) ; Sta Rita Exper Range Florida Cyn Hdqts. 25.VII.2013 at UV & Hg vapor lights (1, BYU) ; Sta Rita Mts. Box Cyn 6.VII.1975 (1, FSCA) ; Santa Rita Mts. Madera Cyn. 30.VII.1973 on Quercus arizonica (1, ERPC) . Pinal Co.: Santa Catarina Mountains 14.V.2022 (1, GBIF) . Santa Cruz Co.: Madera Cyn . 5.IV.1975 (1, FSCA); Patagonia Mts. Finley & Adams Canyon 18.VII.1991 UV Light (1, TAMU). ARKANSAS: Benton Co.: Illinois R. at Osage Ck. dry ice baited sticky trap 26.V.1972 (1, TAMU). FLORIDA: Alachua Co.: Gainesville 29.III-14.IV.1989 (1, FSCA ( Riley et al, 2001)). KANSAS: Kiowa Co.: 13 S Greensburg 4.VI.1995 pack rat [ Neotoma sp. ] midden Salsbury (1, ERPC); Kyowa Co . 6.VI.1923 (2, FIMU). MISSOURI: Dent Co.: Dent Co . 21.VIII.1974 (1, ERPC); Jackson Co.: Jackson Co . 21.VI.1980 (2, ERPC). NEW MEXICO: Hidalgo Co.: Animas Mts. Indian Creek 5.VIII.1979 1737m at light (1, TAMU). Lincoln Co.: Arroyo Seco 5.VIII.2001 (1, BYU); Beaver Dam State Park 1480m 24.VI.2014 (1, BYU). NORD CAROLINA: Durham Co. : Durham Trinity Park 11.VII.2022 (1, GBIF). OKLAHOMA: Latimer Co.: Latimer Co. VI.1987 & VI.1988 & V.1989 & VI.1990 (4, ERPC). TEXAS: Brewster Co.: Green Gulch Big Bend Nartional Park 14.VIII.1968 (1, TAMU). Cameron Co.: Sabal Palm Grove Ref (site 11) 4.IV.2009 (1, TAMU). Hidalgo Co.: Santa Ana Nat. Wdlf Refuge 8.V.1991 (2, TAMU). Jeff Davis Co.: 11 mi W Ford Davis 2.VII.2004 (3, TAMU); Mt. State Park 5.VII.2003 (1, TAMU); Fort Davis 18.VI.1965 (3, FIMU); N of Ft. Davis Hwy 118 16.VI.1963 (1, FIMU). Lubbock Co.: Sunrise Ln. Ransom Canyon 7.V.2022 (1, GBIF). Sabine Co.: 9 Mi E Hemphill 24.IV.1989 (1, TAMU); 9 Mi E Hemphill sift debris from Neotoma nest (2, TAMU). Starr Co.: Falcon Dam 25.IV.1976 (1, TAMU). Uvalde Co.: 3–7 mi SE Concan 11.V.1978 (2, FIMU). UTAH: Garfield Co.: Escalante Ri. Mouth of Calf Creek (1, BYU); Summit Co. : Bellevue 3400 ft 21. III.1919 (1, BYU). Washington Co.: Zion nat. Park Oak Creek 23.VII.1981 (1, BYU) .

Additional data from literature. Baja California Sur ( Horn, 1895; Andrews & Gilbert, 2005); Arizona, Texas, Baja California ( Schaeffer, 1906); Chihuahua, Sonora and Morelos ( Pallister, 1953); Morelos ( Niño-Maldonado et al., 2016); Tamaulipas ( Sánchez-Reyes et al., 2015; Sánchez-Reyes et al., 2016); Baja California Sur, Chihuahua, Guerrero, Morelos, Oaxaca, Sonora, Tamaulipas ( Ordóñez-Reséndiz & López-Pérez, 2021); Arkansas, Florida, Missouri, Oklahoma, Texas ( Riley et al., 2001). “ Riley et al. (2003) also report Maryland and Tennessee, without more precise indication, as part of the species’ range. I did not see any specimens from those states. Suffrian (1866) reported this taxon (under the name of Scolochrus cazicus ) also from “Neu Grenada ”. This would extend the range of the species southward at least up to Panama. However, further data are needed to better outline the southern border of its distribution.

Distribution. El Salvador, Guatemala, Mexico, Panama?, U.S.A.

Diagnosis. The species is highly variable in colouration. However, the chromatic pattern is always very distinctive and allows to easily separate this species from the closest ones. The shape of the aedeagal median lobe leads to suppose that the closest species is G. lecontii .

Description of male. BL = 3.9–5.0 mm, BW = 2.5–3.1 mm, PL = 1.4–2.0 mm, PW = 1.9–2.9 mm. Interocular distance 0.0–2.6 % of BL.

Head ( Fig. 9d ) white with lower part of frontoclypeal area, from antennal insertion to lower rim, black. Ocular lines and narrow space between upper lobes of eyes (when present) black as well. Labrum white. Vertex bright, very shallowly punctured with short setae. Surface of frontoclypeal area bright as well, with shallow punctation and scattered short setae above all on lower part of frontoclypeus. White patches mildly raised with respect to black area. Mid-cranial suture short but detectable on anterior part of vertex and between ocular upper lobes. Upper lobes of eyes almost in contact along midline, but very narrow space in between often discernible. Ocular lines narrow, but fairly detectable because black against light background. Ocular canthus large, not differentiated in punctation and setosity from the remainder of frontoclypeal surface. Antennae ( Fig. 9n ) rather short, brown with antennomeres 3–5 only slightly paler than the subsequent ones, bright, subcylindrical; antennomeres 6–11, dull, more flattened and more diffusedly setose.

Pronotum red with whitish anterior margin. Large, obtusely subtriangular, black band extended from anterior white margin to middle of disc. Pronotal shape roughly elliptical, scarcely transverse, regularly convex on disc. Lateral margins narrow, not visible from above, regularly curved so that maximum width nearly at middle. Surface moderately shiny usually with close, well-impressed punctation almost evenly distributed, as size and closeness, across whole surface. Often punctures coarse and coalescent to look like transverse rugosity. Posterolateral impressions weak but detectable.

Scutellum whitish with black apical rim to almost totally brownish, subtriangular with apex shortly truncated. Surface minutely punctured, with few, short setae.

Elytron black with white stripe along suture and apical margin. Further white marking along basal margin (externally to fourth row of punctures) and anterior third of lateral margin. Short further stripe between anterior sections of eighth and ninth rows. Epipleuron white. Sometimes elytron white on anterior half, black on posterior one (described as form “ suffriani ”) ( Fig. 9f ). In other cases, elytron black with white patch surrounding scutellum and extended along suture up to midline; this white patch often laterally reaching second to fourth row of punctures (described as form “ cazicus ”) ( Fig. 9e ). Again, at times whole surface of elytra totally black. In this latter case pronotum red with black anterior margin ( Fig. 9g ). Elytral outline short with sides almost straight and convergent posteriorly. Lateral margins narrow, simultaneously visible from above only along posterior half. Elytral surface barely flattened on disc, moderately shiny, with strong punctation arranged in regular rows, well visible up to posterior clivus. Intervals flat. Postscutellar area fairly raised. Humeral callus prominent, impunctate. Epipleuron smooth, impunctate, with convex surface.

Pygidium white with margins narrowly black, very rarely black pattern more extended until covering whole surface. Surface matt, covered with close shallow punctures and appressed, pale setae.

Ventral parts of thorax black with posterior half of hypomera red. Abdominal ventrites 1–4 black with white lateral margins. Fifth ventral ventrite white. Medial projection of first abdominal ventrite white. Often white colour extended to almost entire ventral surface. Hypomera, mesepimera and mesepisterna shiny, with well-impressed, sparse punctures and few, scattered setae. Remainder of ventral surface of thorax and abdomen covered with fine punctures and regularly distributed, thick, whitish setae. Prosternal process large, with sides almost straight between anterior coxae, then converging in large, round apex; surface slightly depressed, uneven, covered with coarse, shallow punctures and sparse, long, semi-erect setae. Legs black or brownish, tinged with white on femora and tibiae.

Median depression on fifth abdominal ventrite very shallow and hardly detectable, barely differentiated in punctation and setosity from remainder of ventrite surface. Ventrite posterior margin straight. Median lobe of aedeagus ( Fig. 9h–m ) with apex rather long, triangular, terminated with blunt median denticle. Aedeagal shaft strongly flattened dorsoventrally, with apex almost straight in lateral view. Ventral surface smooth, with a faint hint of longitudinal median carina. Setose depressions reduced to tufts of curved setae along base of aedeagal apex.

Female. Habitus in Fig. 9a–b (HT). BL = 4.3–5.8 mm, BW = 2.7–3.7 mm, PL = 1.7–2.1 mm, PW = 2.5–3.3 mm. Interocular distance 7.0–10.3 % of BL.

Females are larger with eyes ( Fig. 9c ) smaller and well separated along the midline. The head surface is matt, covered with coarse, shallow punctures and scattered setosity. The head is black with a transverse white band between the ocular canthi. Irregular white patches at times on vertex as well.

The fifth abdominal ventrite in females has a large, rounded and deep pit. The bottom of the pit is black (while the remainder of the ventrite is yellow), shiny and covered with small punctures and very short setae. The vasculum of the spermatheca ( Fig. 9p ) is scarcely pigmented, sickle-shaped with fairly swollen proximal lobe. The distal lobe is short, slender, tapered with a rather acute apex fairly bent downwards. The ampulla is not pigmented, moderately lengthened. The duct insertion on the ampulla is is simple, very short and not pigmented. The sperm gland insertion is longer and bent upwards. The duct is uniform in size, slender, not coiled, devoid of turns or sometimes with a single turn close to the vasculum. The insertion on the bursa copulatrix is simple, neither swollen nor pigmented.

Remarks. This taxon is very variable, not only in the colour pattern, but also in the size, the dorsal punctation, and even in the body outline (compare for instance fig 9e and 9f, where the difference is actually quite surprising). The distance between upper lobes of the eyes in male specimens can also be rather different. Nonetheless, it was not possible to detect any regularity in the variability that could not be interpreted as individual variation and therefore justify the maintenance or creation of distinct taxonomic entities. Quite interesting is also the chromatic peculiarity of the species, whereby the light colour, when present, is not yellow, as it happens, to the best of my knowledge, in all the other species of the same genus, but frankly white (even though it tends towards light yellow in dried specimens), contributing to give G. montezuma a truly unique appearance even compared to similar species.

Figures

FIGURE 9. G. montezuma. a–b: habitus, dorsal and lateral view (HT); c: frontal view, female (HT); d: frontal view, male (Guayamas, Mexico); e–g: colour variations (e: male, Cacahuamilpa, Mexico; f: female, Jalapa del Marques, Mexico; g: female, Sabine Co., Texas); h–m: median lobe of aedeagus in ventral (h), dorsal (k) and lateral (m) view [0.5 mm]; n: antenna [0.9 mm]; p: spermatheca [0.3 mm]. hd: setose depressions. Square brackets: length of scale segment.

FIGURE 13. Distribution maps for the Griburius scutellaris species-group. a: G. lecontii, Mexican range; b: G. lecontii, U. S. range; c: G. montezuma, Mexican and Central American range; d: G. montezuma, U. S. range; e: G. rileyi; f: G. scutellaris. Black markings: data from specimens analyzed during the study. Red markings: data from the public database Global Biodiversity Information Facility (GBIF.com) or literature data. Black stars: the presence in the state is reported in the consulted literature, but without more precise data on the location.