TRANSACTIONS RESEARCH ARTICLE TAES 140: 209-236 AMERICAN ISSO 0002-8320 ENTOMOLOGICAL http://taes.entomology-aes.org/ SOCIETY
Revision of the Mexican genus Philocleon (Orthoptera: Acrididae: Melanoplinae)
Daniel Otte
Academy of Natural Sciences, Philadelphia, PA 19103 email: [email protected]
ABSTRACT
The grasshopper genus Philocleon, with six previously known species (nigrovittatus (Stal), spatulatus Roberts, anomalus Roberts, luceroae Fontana and Buzzetti, scudderi (Hebard), and ottei Fontana & Buzzetti), is revised to include the following eight new species: zygon, zima, illa, cledon, axiton, azumai, iropon, and erissa. The subspecies nigrovittatus spatulatus Roberts is raised to species status. The fourteen species are placed into the four following species groups: Anomalus group, Illa group, Nigrovittatus group, and Scudderi group.
[Key Words: Acridoidea, Acrididae, Melanoplinae, Melanoplini, Philocleon, Mexico, new species]
INTRODUCTION
The genus Philocleon Scudder 1897 is known Derivation of Names only from Mexico and is distributed from Coahuila Philocleon azumai is named in honor of Don and Nuevo Leon in the north to Guerrero and Azuma who served as collection manager in the Oaxaca in the south. Prior to the present study five Entomology Department of the Academy of Natural species were known: P. nigrovittatus (Stål 1875), Sciences for many years. All other names of new P. scudderi (Hebard 1932), P. anomalus Roberts species are random combinations of letters with no 1941, P. luceroae Fontana and Buzzetti 2007, and known meanings. P. ottei Fontana and Buzzetti 2007. A subspecies of nigrovittatus (Stål), P. nigrovittatus spatulatus SYSTEMATICS Roberts 1947 is here raised to species level. In this paper we recognize four species groups and add Diagnosis of Genus eight new species: P. zygon n. sp., P. illa n. sp., P. zima n. sp., P. cledon n. sp., P. axiton n. sp., P. 1. Cerci thick at base, narrowing strongly after first azumai n. sp., P. iropon n. sp., and P. erissa n. sp.. third, and strongly incurved after first half (Fig. 17). PHILOCLEON Scudder 1897 2. Furculae present, short, closely spaced, and situated in a median furrow (Figs. 17). Type species Pezotettix nigrovittatus Stal (now Philocleon 3. Aedeagus narrow and pointed (Figs. 5, 7, 9, 10, nigrovittatus nigrovittatus), by original monotypy. 12, 13). Scudder. 1897. Proceedings of the American Academy of 4. Often with brilliant yellow, green, orange, red, Arts and Sciences 32: 203, 206 >> Philocleon and black coloration (Figs. 1-4). Hebard. 1932. Transactions of the American Entomological Society 58(3): 293 >> diagnosis, type Relationship to other Melanoplinae remains species designation Roberts. 1947. Proceedings of the Academy of Natural unknown. Other genera are currently under study, Sciences Philadelphia 99: 226 >> Melanoplinae but because the subfamily and the tribe Melanoplini Fontana and Buzzetti. 2007. Atti Acc. Rov. Agiati 8 7(B): is large much work needs to be completed before 105 >> Philocleon relationships can be determined.
Publication date: 15 May 2014 urn:lsid:zoobank.org:pub:731481BE-035F-4FCE-9DC9-A86FB5AB07C2 210 REVISION OF PHILOCLEON
Philocleon consists of four distinct species Nigrovittatus Group (southern Mexico)— groups, based on morphology, as follows: Anomalus nigrovittatus and spatulatus Group with two species: P. anomalus Roberts and P. zygon; Illa Group with five species: P.zima , P. illa, 1. Males and females with small elongate or oval P. cledon, P. axiton, and P. azumai; Nigrovittatus forewings, these covering auditory tympana. Group with two species: P. nigrovittatus (Stål) 2. Hind femur banded longitudinally with and P. spatulatus (Roberts); and Scudderi Group red, black and ivory; chevron dark; lower with five species: P.scudderi (Hebard), P. luceroae marginal areas black or black and red. Fontana and Buzzetti, P. ottei Fontana and Buzzetti, 3. Upper dark band of pronotum usually extending P. iropon n. sp. and P. erissa n. sp. from anterior to posterior margin. 4. Upper face of metazona of pronotum orange to IDENTIFICATION OF SPECIES GROUPS red. 5. Antenna yellow green or orange. (Items in bold face are especially useful in 6. Upper posterior corner of lateral lobes of separating the species groups) pronotum without a distinct pale triangle.
Anomalus Group (southern Mexico) — anomalus Scudderi Group (central and northeastern Mexico) and zygon — scudderi, luceroae, ottei, iropon, and erissa
1. Males and females with small elongate or oval 1. Males and females without any forewings and forewings; these cover the auditory tympana. without auditory tympana. 2. Hind femur green, yellow green or orange, 2. Hind femur with a row of dark markings without dark markings anterior to knees. along lower marginal area; chevron of 3. Upper dark band on pronotum extends from medial area usually very distinct, either anterior to posterior margin. green or black. 4. Upper face of metazona of pronotum green, 3. Upper dark band of pronotum usually extending never with orange, or reddish. onto metazona (and to posterior margin). 5. Antennae never yellow, orange or reddish. 4. Upper face of metazona of pronotum orange to 6. Upper posterior corner of lateral lobes of red. pronotum without a distinct pale triangle. 5. Antenna yellow green or orange. 6. Upper posterior corner of lateral lobes Illa Group (southern Mexico) — zima, illa, cledon, of pronotum always with a distinct pale axiton, and azumai triangle.
1. Males and females with small elongate or oval forewings, these covering auditory tympana. KEY TO SPECIES GROUPS AND SPECIES 2. Hind femur yellow or orange, with a single strong dark area just anterior to narrowest 1 Males and females with small forewings, and point of femur. with auditory tympana ...... 2 3. Upper dark band of pronotum usually extending — Males and females without forewings and from anterior to posterior margin. without auditory tympana—Scudderi Group . . . 4. Upper face of metazona of pronotum orange to ...... 10 red. 5. Antenna yellow, green, or orange. 2 Hind femora without dark marking anterior to 6. Upper posterior corner of lateral lobes of black knees; metazona without red or orange pronotum without a distinct pale triangle. coloration (Figs 2A, 2B) —Anomalus Group ...... 3 DANIEL OTTE 211
— Hind femora with dark marks anterior to black 9 Upper dark band of pronotum containing pale knees; metazona red or orange (Figs 2C-G, 3, 4) areas in males, variable in females (Figs 3F-H) ...... 4 ...... spatulatus — Upper dark band of pronotum solid, sometimes 3 Male genitalia as in Figs 5A-C, 6A-E (Guerrero containing pale areas in females (Figs 3A-E) and Morelos) ...... anomalus ...... nigrovittatus — Male genitalia as in Figs 5D, 6F, G (Oaxaca) ...... zygon 10 Chevron of hind femur greenish. Without a dark band at end of chevron (Figs 4G, H, I). 4 Hind femora with a single dark mark near Aedeagus elongate (Figs 12A-E) . . . . 11 narrowest part of femur (Fig. 2) —Illa Group — Chevron of hind femur very dark in males (Figs ...... 5 4A-D). With a dark band at end of chevron. — Hind femora with widespread dark markings Aedeagus short (Figs 13A-D) ...... 12 along femur, black along lower marginal area (Figs 1, 3) —Nigrovittatus Group ...... 9 11 Lower end of lateral lobe of pronotum with a dark band extending to margin (Figs 4A, B) . . . 5 Hind femora with a row of dark spots on ...... luceroae lower marginal area and a red streak on upper — Lower end of lateral lobe of pronotum without a marginal area; and dark spot adjacent to dark band extending to margin (Fig. 4H) . . narrowest part indistinct (Fig. 2C) (Colima) ...... ottei ...... zima — Hind femora without a row of dark spots on 12 Aedeagus very short (Figs 13A-D) ...... 13 lower marginal area; with a distinct dark mark — Aedeagus intermediate in length (Fig. 12E) adjacent to narrowest point (Figs 2D, E, F, G) (Coahuila, Nuevo Leon) ...... iropon ...... 6 13 Crest of pronotum strongly arched (Fig. 4F) 6 Pronotum strongly arched; hind femur with an (Coahuila) ...... erissa ivory ring at narrowest part, preceded by a broad — Crest of pronotum not strongly arched (Figs 4C, black band (Figs 1, 2E) (Nayarit) . . . . azumai D, E, G) (San Luis Potosi, Zacatecas, Durango) — Pronotum not strongly arched; femora not ...... scudderi marked as above (Figs 2D, F, G) ...... 7
7 Hind femur with scattered dark markings in chevron of medial area (Fig. 2F) (near coast of Jalisco) ...... axiton — Hind femur without scattered dark markings on chevron, but with a distinct dark spot adjacent to narrowest part (Figs 2D, E, G) . . . 8
8 Upper dark band of pronotum containing lighter markings and not extending onto metazona (Fig 2D) (Coastal Jalisco) ...... cledon — Upper dark band solid, and usually extending onto metazona (Fig. 2G) (Coastal Jalisco, and Colima ...... illa 212 REVISION OF PHILOCLEON
GROUP 1: Anomalus GROUP on Hwy 190, 15 Sept 1959, 5750 feet (Cantrall and Cohn #127)—3M 2 F (Specimens H12, PH- 1. Philocleon anomalus Roberts 1941 16) (UMMZ). — MICHOACAN: Morelia, 6-8000 Figures 1, 2A, 5A-C, 6A-E, 15A, 17 feet, 4 Nov 1938 (Roberts)—6M 3F (ANSP). — GUERRERO: Mazatlan, 5000 feet, 14 Nov 1938 Philocleon anomalus Roberts. 1941. Notulae (Roberts) 1M 1F (ANSP). Naturae 76: 1. Holotype male, Mexico, Guerrero, 16 miles south of Chilpancingo, 2. Philocleon zygon new species Academy of Natural Sciences, Philadelphia. urn:lsid:zoobank.org:act:E6DB6BA2-0F7B-4BAB-B36E- Philocleon anomalus. García, García and Fontana. 54B872866A96 2006. Entomologia Mexicana 5(2): 250. Figures 2B, 5D, 6F, G, 15B, 17 Philocleon anomalus. Fontana, Buzzetti and Mariño-Pérez. 2008. Chapulines, Langostas, Holotype male.— Mexico, Oaxaca, 1.7 miles Grillos y Esperanzas de México. Guía SE of El Camaron (42 miles NW of Tequisistlan fotográfica - Grasshoppers, Locusts, Crickets on Hwy 190), 14 Sept 1059, 2000 feet (Cantrall & Katydids of Mexico. Photographic guide & Cohn #118)—Specimen H15, University of 190-191 Michigan Museum of Zoology. Philocleon anomalus. García García, Fontana, Recognition.— Head: Antennae color pale Marini, Equihua-Martiniez, Sanchez-Escudero, brown or orange brown. Face color uniform grey- Valdez-Carrasco & Cano-Santana. 2008. Jour. brown. Pronotum: Lateral lobe color: upper two Orth. Res. 17(1):85 >> Note: Found in cactus thirds dark brown. Pronotal disk color: interrupted scrub habitat. pale band along margins, median half pale to dark brown. Forewings present, slightly longer than half Distribution.— Guerrero, Michoacan, Pueblo. the pronotal length. Cerci strongly bent at middle. Recognition.— Head: Antennae color, pale Furculae very small, globular. Front legs green. brown. Face color greenish. Pronotum: Lateral lobe Middle legs green. Hind legs entirely green or color: upper half dark brown, lower half green. yellow green, with entirely black knees. Tibia color Pronotal disk color entirely green. Forewings bluish. present, slightly longer than half length of Body measurements (mm).— Body Length: 18- pronotum. Cerci strongly bent, flattening at apex, 20 (males), 22-24 (females). Hind femur length: 11- sometimes turning black. Furculae short, blunt, as 12 (males), 13 (females). long as wide. Front legs yellow green, with green Specimens examined.— OAXACA: 45 miles tibiae. Middle legs yellow green, with green tibiae. NW of Tequisistlan (1 mile S of El Camaron), Hind legs uniform pale green, with entirely black 21 Dec 1958, 2000 feet (TJ Cohn #384)—2M 1F knees. Tibia color blue green. (Specimen H14) (UMMZ). — 1.7 miles SE of El Body measurements (mm).— Body Length: 15- Camaron (42 miles NW of Tequisistlan on Hwy 16 (males), 17-23 (females). Hind femur length: 10- 190), 14 Sept 1059, 2000 feet (Cantrall & Cohn 10.5 (males), 10.5-13 (females). #118)—4M 2F (Specimens H15, PH-11) (UMMZ). Specimens examined.— MEXICO: GUERRERO: 16 miles S of Chilpancingo, 4000 feet, 14 Sep 1940 (Bolivar & Roberts)—10M 4F PARATYPES (ANSP). — 16 road miles S of Chilpancingo (km298 on Hwy 95) 3800 feet (Cantrall and Cohn #147)—3M 2F (Specimens PH- 15, H9) (UMMZ). — 9 road miles NE of Taxco, 1.7 road miles SW of Acuitlapan, 5700 feet, 16 Sept. 1959 (Cantrall & Cohn #136) (Specimen H10) (UMMZ). — PUEBLO: 6 miles NW of Petlalcingo, DANIEL OTTE 213
GROUP 2: Illa GROUP longer than wide. Front legs yellow-orange with green tibiae. Middle legs same as front legs. Hind 3. Philocleon zima new species legs: Medial area yellow-green, with a dark band urn:lsid:zoobank.org:act:E6DB6BA2-0F7B-4BAB-B36E- and a pale band anterior to the black knee; lower 54B872866A96 posterior quarter of knee pale. Upper marginal area Figures 2C, 7B, 8B, 15D, 17 yellow-green. Lower marginal area yellow-green. Tibia color bluish. Holotype male.— Mexico: Colima, 8 miles Body measurements (mm).— Body Length: 19- SW of Colima, 28 Nov 1958 (TJ Cohn #325) — 19 (males), 26-27 (females). Hind femur length: ca. Specimen PH-4, University of Michigan Museum 12 (males), ca. 14 (females). of Zoology. Specimens examined.— MEXICO: COLIMA: Body measurements (mm).— Body Length: 26. 11.3 miles SW of Colima (on Hwy 110), 28 Sept Hind femur length: 14. 1959, 1300 feet (Cantrall & Cohn #192)—4M 2F Recognition.— Head: Antennae color orange. (Specimens H11, H30, PH-3) (UMMZ). — 13 Face color greenish. Pronotum: Lateral lobe color miles S of Colima, 28 Nov 1958, 1000 feet (TJ mostly dark in upper two thirds, this band present Cohn #324) (UMMZ). — JALISCO: 18.3 miles N only in anterior two thirds. Pronotal disk color with of Barra de Navidad (on Hwy 80) 2 Sep 1968 (TJ dark brown band in anterior two thirds, slightly Cohn #32) (UMMZ). — 16.2 miles N of Barra de orange in posterior third. Forewings present, about Navidad (on Hwy 80) 2 Sep 1968 (TJ Cohn #33) as long as half pronotal length. Cerci gradually (Specimen H8) (UMMZ). rounding towards median line. Furculae slender and pointed. Front legs greenish brown. Middle legs 5. Philocleon cledon new species green brown. Hind legs: Medial area greenish in urn:lsid:zoobank.org:act:8E2046FD-C357-4A20-94A6- upper half, yellowish in lower half. Upper marginal ED15501B1464 area reddish. Lower marginal area reddish, with a Figures 2D, 7C, 8C, 15E, 17 row of black dots. Tibia color greenish-yellow Specimens examined.— COLIMA: 8 miles SW Holotype male.— Mexico: Jalisco, 16.2 miles of Colima, 28 Nov 1958 (TJ Cohn #325) —1M N of Barra de Navidad (on Hwy 80) 5 Oct 1970 (PH-4) (Specimen PH-4) (UMMZ). (Cohn and Cohn #44)— Specimen H41. University of Michigan Museum of Zoology. 4. Philocleon illa new species Recognition.— Head: Antennae color orange. urn:lsid:zoobank.org:act:39C54A33-7622-4918-A286- Face color greenish. Pronotum: Lateral lobe color B04EA79AFA9C with black upper two thirds, this black band located Figures 2G, 7A, 8A, 15C, 17 in front two thirds, grooves black. Pronotal disk color: median dark band in front two thirds, yellow Holotype male.— Mexico: Colima, 11.3 miles orange in posterior third. Forewings present, about SW of Colima (on Hwy 110), 28 Sept 1959, 1300 half as long as pronotum. Cerci gently incurved. feet (Cantrall & Cohn #192)— Specimen H11. Furculae thick and short. Front legs yellow-green, University of Michigan Museum of Zoology. tibiae more green. Middle legs yellow-green, tibiae Recognition.— Head: Antennae color orange- green. Hind legs: Medial area greenish. Upper brown. Face color greenish. Pronotum: Lateral marginal area greenish. Lower marginal area lobe color: upper two thirds black or dark brown, orange. Tibia color bluish. sometimes containing paler islands, groves black. Body measurements (mm).— Body Length: 22- Pronotal disk color with pale lateral margins, 23 (males), 22-28 (females). Hind femur length: 13- median band black in anterior two thirds, but with 13.5 (males), 14-15 (females). a narrow median pale stripe, orange in last third. Specimens examined.— MEXICO: JALISCO: Forewings present, slighly longer than half pronotal 16.2 miles N of Barra de Navidad (on Hwy 80) 5 length. Cerci strongly bent at middle poin. Furculae Oct 1970 (Cohn and Cohn #44)—3M 2F (Specimen 214 REVISION OF PHILOCLEON
H41) (UMMZ). — 18.3 miles N of Barro de Recognition.— Head: Antennae color orange Navidad (on Hwy 80) 5 Oct 1970 (Cohn and Cohn brown. Face color orange. Pronotum: Lateral #42)—1M 1F (Specimen H7) (UMMZ). — 12.4 lobe color upper half black in anterior two thirds, miles N of Barro de Navidad (on Hwy 80) 5 Oct grooves black. Pronotal disk with a tall median 1970 (Cohn and Cohn #43)—1M (UMMZ). crest, black in front two thirds, orange in last third. Forewings about half as long as pronotum. Cerci 6. Philocleon axiton new species gently incurved. Furculae short, pointed. Front legs: urn:lsid:zoobank.org:act:B1319655-4E67-4AC8-848A- femur black on outer face, tibia pale except black at 8812448C7682 base. Middle legs: femur pale brown, black at distal Figures 2F, 7D, 8D, E, 15F, 17 end, tibia black at both ends. Hind legs: Medial area orange, darkening to black in last quarter. Upper Holotype male.— Mexico: Jalisco, 6.2 miles marginal area orange. Lower marginal area orange- SW of Autlan Plaza, 5 Oct 1970 (Cohn and Cohn red. Tibia color blue-brown. #41)— Specimen H42. University of Michigan Body measurements (mm).— Body Length: ca Museum of Zoology. 22 (males), 28-32 (females). Hind femur length: ca Recognition.— Head: Antennae color orange. 13 (males), 28-32 (females). Face color yellow-green. Pronotum: Lateral lobe Specimens examined.— MEXICO: NAYARIT: color: dark band in upper half interrupted, grooves 35.4 miles SW of Las Varas (in hills), 8 Oct 1970 black. Pronotal disk color: with median dark band (Cohn and Cohn #41)—3M 1F (Specimens H45, in anterior two thirds, orange in posterior third. PH-12, PH-12, PH-13, PH-14F) (UMMZ). Forewings: about two thirds length of pronotum, with green margins. Cerci gently incurved, very GROUP 3: Nigrovittatus GROUP thick at base. Furculae very short and thick. Front legs yellow green. Middle legs yellow green. Hind 8. Philocleon nigrovittatus (Stål) legs: Medial area yellowish with black median Figures 1, 3A-D, 9A-F, 11A-F, 15G, 17 markings through most of its length. Upper marginal area with orange band. Lower marginal Pezotettix nigrovittatus Stål 1875. Bihang Kungl area yellowish or orange. Tibia color blue-green. Svenska Vet. Akad. Handl 3(14): 32. Type Body measurements (mm).— Body Length: ca. male, Mexico, NHRS Stockholm. 22 (males), ca. 26 females). Hind femur length: ca. Pezotettix nigrovittatus. Scudder, S.H. 1897. Proc. 13 (males), ca. 17 (females). Amer. Acad. Arts and Sci. 32:206. Specimens examined.— MEXICO: JALISCO: Pezotettix nigrovittatus. Hebard. 1932. Trans. Amer. 6.2 miles SW of Autlan Plaza, 5 Oct 1970 (Cohn Entomol. Soc. 58(3):293 >> Note: Distribution and Cohn #41)—3M 1F (Specimens H42, PH-18) (UMMZ). Recognition.— Head: Antennae color reddish brown, not orange. Face color greenish-brown. 7. Philocleon azumai new species Pronotum: Pronotal disk color median dark band urn:lsid:zoobank.org:act:B98C0B2E-88E2-4E4B-A6B5- in anterior two thirds, red in last third. Forewings 4C564FFEB339 brown with a black central band. Furculae long, Figures 1, 2E, 7E, 8F, G, 15I, 17 slender widely separated. Hind legs: Upper marginal area reddish. Lower marginal area black, Holotype male.— Mexico: Nayarit: 35.4 miles red in lower third. SW of Las Varas (in hills), 8 Oct 1970 (Cohn Body measurements (mm).— Body Length: and Cohn #41)— Specimen H45, University of 17-23 (males), 24-27 (females). Hind femur length: Michigan Museum of Zoology. 11.5-14 (males), 12-14 (females). Etymology.— This species is named honor of Don Azuma who served as Collection Manager in Specimens examined.— MEXICO: JALISCO: the Department of Entomology for many years. Barranca de Oblatos, 7 miles N of Guadalajara, DANIEL OTTE 215
1 Oct 1959, 4850 feet (TJ Cohn #201)—4M 2M 9. Philocleon spatulatus Roberts, new status (Specimens G19, PH-28) (UMMZ). — JALISCO: Figures 3F-H, 10A-C, 11G-M, 15H, 17 Tequila, 4300 feet, 12 Sep 1938 (Roberts)—5M 3F (ANSP). — MICHOACAN: 3.5 miles W of Philocleon nigrovittatus spatulatus Roberts. 1947. Jiquilpan, on Hwy 110, 28 Sep 1959, 6150 feet Proc. Acad. Nat. Sci. Philad. 99:227. Holotype (Cantrall and Cohn #187)—2M 1F (Specimens male, Mexico, Michoacan, Morelia, 4 ix 1938. H18) (UMMZ). — 0.3 miles S of Tzitzio, on Hwy Academy of Natural Sciences Philadelphia. 49, 24 Sep 1959, 4959 feet (Cantrall and Cohn Philocleon nigrovittatus spatulatus. Fontana, #168)—2M 2F (UMMZ). — 3 miles SE of Tequila, Buzzetti & Mariño-Pérez. 2008. Chapulines, 22 Nov 1958, 4100 feet (TJ Cohn #307)—1M Langostas, Grillos y Esperanzas de México. (Specimen PH-30) (UMMZ). — 4.5 road miles Guía fotográfica - Grasshoppers, Locusts, W of Jiquilpan, 1 Dec 1958, 6100 feet (TJ Cohn Crickets & Katydids of Mexico. Photographic #335)—2M (Specimen H32) (UMMZ). — guide 193-194. Cojumatlan, Lake Chapala, 4-5500 feet, 8 Sep 1938 (Roberts)—6M (ANSP). — JALISCO: 14.3 road Body measurements (mm).— Body Length: ca. miles S of Ixtlahuacan del Rio (1.3 road miles S of 24 (males). Hind femur length: ca. 14 (males). Rio Verde Bridge) 3380 feet, 1 Nov 1974 (Cohn and Recognition.— Head: Antennae color orange. Cohn #126)—2M (UMMZ). — NE Guadalajara Face color brownish. Pronotum: Lateral lobe color (Barranca de los Oblatos) 12 Nov 1974 (Cohn and upper half with black band, with islands of brown, Cohn #126)—5M (UMMZ). — Colotitlan, 4500 grooves black. Pronotal disk color: median dark feet, 19 Aug 1940 (Roberts)—13M 8F (ANSP). band in front two thirds, last third red. Forewings — GUERERO: 9 road miles NE of Taxco, 1.7 road mostly pale. Front legs: femur black on outer face, miles SW of Acuitlapan, 5700 feet, 17 Sep 1959 tibiae greenish. Middle legs: femur black on outer (Cantrall and Cohn #137)—2M 2F (Specimen H31) face, tibiae greenish. Hind legs: medial area upper (UMMZ). — MICHOACAN: 7 miles N of Tzitzio half with black concentrated in two bands. Upper (on Hwy 49) 24 Sep 1959, 6400 feet (Cantrall and marginal area dark red. Lower marginal area black Cohn #169)—2M 2F (Specimen H16) (UMMZ). in upper half, red in lower half. Tibia color mostly — 4.2 miles E of Morelia (on Hwy 15) 24 Sep 1959 black, brown in central area (Cantrall and Cohn #170)—4M 2F (Specimen H17) Specimens examined.— MEXICO: JALISCO: (UMMZ). — COLIMA: Vulcano (L. Conrad)—1M 0.4 miles S of Huentitlan del Bajo (5 miles N of 1F (UMMZ). — MORELOS: Tepoztlan, 5000 Guadalajara, 30 Sep 1959, 5100 feet (TJ Cohn feet, 28 Aug 1938 (Roberts)—5M 1F (ANSP). — #199)—2M 2F (Specimens G23, PH-29) (UMMZ). JALISCO: 1 mile S of Pihuamo, 26 Nov 1958, 2850 — 15 miles W of Zapotlanejo, 23 Nov 1958, feet (TJ Cohn # 314)—2M 1F (Specimens H29, 5200 feet (TJ Cohn #308)—1M (Specimen H27) PH-21) (UMMZ). — MORELIA: Cuernavaca, 4500 (UMMZ). — 10.5 miles NW of Leon (town center) feet, 1 Sep 1936 (Roberts)—12M 11F (ANSP). (ridge summit) 11 Nov 1970 (Cohn and Cohn #87)—1M (Specimen H43) (UMMZ). — 13 mile SW of Guadalajara, 4 miles NW of Santa Cruz, 24 Nov 1958, 5300 feet (TJ Cohn #310)—4M 1F (UMMZ). — 26 miles W of Guadalajara, 4500 feet, 11 Sep 1938 (Roberts)—6M (ANSP). — Tequila, 4300 feet, 12 Nov 1938 (Roberts)—1F (ANSP). — MICHOACAN: Morelia, 6-8000 feet, 4 Sep 1938 (Roberts)—6M 1F (ANSP). — Zamora, 5500 feet, 8 Sep 1938 (Roberts) (ANSP). — Lake Patzcuaro, 6500 feet, 4 Sep 1938 (Roberts)—1M 1F (ANSP). — between Guadalajara and Chapala, 8 Sep 1938 (Roberts) (ANSP). — GUERRERO: km 234, 216 REVISION OF PHILOCLEON
between Iguala and Rio Balsas, 2000 feet, 13 Sep (males), 22-26 (females). Hind femur length: 8-11 1940 (Bolivar and Roberts)—1M (ANSP). — 22 (males), 10-14 (females). road miles NE of Arcelia, 4000 feet, 8 Dec 1958 (TJ Specimens examined.— ZACATECAS: Cohn #362)—2M 1F (Specimen PH-31) (UMMZ). Camacho, Nov 1897 Holotype M, 3 paratype — MICHOACAN: 23 miles E of Nueva Italia (on males, 4 paratype females (ANSP). — 41.6 miles Huacana road) 25 Sep 1959, 950 feet (Cantrall and SE Sombrerete, Hwy 45, 11.8 miles SE of El Sauz, Cohn #175)—1M (Specimen H13) (UMMZ). 7200 feet, 25 Oct 1974 (Cohn and Cohn #95)—8M 6F (Specimens PH-32, PH-36) (UMMZ). — 5.6 GROUP 4: Scudderi GROUP miles SW of Sombrerete, west of road junction on Hwy 45, 7850 feet, 25 Oct 1974 (Cohn and Cohn 10. Philocleon scudderi (Hebard) #96)—2F (UMMZ). — 14.8 miles NE of Nieves, Figures 1, 4C, E, D, G, 13A-C, 14I, J, K, 16D, 17 42.8 miles SW of Camacho, 10 Nov 1970 (Cohn and Cohn #85)—1M (Specimen PH-9) (UMMZ). Philocleon nigrovittatus Scudder, S.H. 1897. Proc. — 33 miles SE of Sombrerete, on Hwy 45, 3.3 U.S. Nation. Mus. 20 (1124):396. Holotype miles SE of El Sauz, 7000 feet, 31 Oct 1974 (Cohn male, Mexico, Camacho. Academy of Natural and Cohn #113)—3M (Specimen PH-37) (UMMZ). Sciences, Philadelphia. — 1.2 miles W of Camacho, 1700 feet, 10 Nov Zacatacris scudderi Hebard. 1932. Trans. Amer. 1970 (Cohn and Cohn #84)—3M 2F (Specimen Entomol. Soc. 58(3):295. Holotype male, PH-38) (UMMZ). — DURANGO: 17.4 miles N Mexico, Camacho. New name for nigrovittatus of Rodeo church, 2.3 miles SW of Alamillo, 1470 Scudder which is a junior homonym of meters, 7 Nov 1970 (Cohn and Cohn #78)—1M nigrovittatus Stal. Academy of Natural 2F (Specimen H2) (UMMZ). — 34.8 miles E Sciences, Philadelphia. of La Zarca on Hwy 30, 30 Nov 1972 (JC LEE Philocleon scudderi. Roberts. 1947. Proc. Acad. #43)—5M (Specimens BB1, BB2, BB4) (UMMZ). Nat. Sci. Philad. 99:226. — Las Mangas, 10.4 miles SW of Durango, on Philocleon scudderi. Fontana & Buzzetti. 2007. Atti Hwy 40, 8 Nov 1970 (Cohn and Cohn #81A)—1M Acc. Rov. Agiati 8 7(B):107. 2F (Specimen PH-42) (UMMZ). — SAN LUIS Philocleon scudderi. Fontana, Buzzetti & Mariño- POTOSI: 38 miles NE of Villa Hidalgo, 4700 feet, Pérez. 2008. Chapulines, Langostas, Grillos 20 Aug 1959 (Cohn and Cantrall #7)—2M 1F y Esperanzas de México. Guía fotográfica - (Specimen PH-39) (UMMZ). — 10 miles NW of Grasshoppers, Locusts, Crickets & Katydids of El Tepeyac, 37 miles NW of Ciudad del Maiz, 3400 Mexico. Photographic guide 195. feet, 22 Aug 1959 (Cantrall and Cohn #17)—2M (Specimen PH-40) (UMMZ). — 12.5 miles east Recognition.— Head: Antennae color orange. of Rio Verde on Hwy 70, 20 Nov 1970 (Cohn and Face color yellowish (green in some females), Cohn #102)—3M 2F (Specimen PH-41) (UMMZ). black around antennae. Without forewings. Without — Nunez, 22 miles SE of Villa Hidalgo, 4900 feet, auditory tympana. Lateral pronotal lobes distinctly 20 Aug 1959 (Cohn and Cantrall #6)—3M 3F (H21, marked with yellow and ivory spots. Upper dark PH-8) (UMMZ). — 17 miles NE Villa Hidalgo, band continuous or interrupted. Front and middle 4950 feet, 20 Aug 1959 (Cohn and Cantrall #5)—1F femora usually orange. Hind legs: medial area (UMMZ). — 16 miles NE of San Luis Potosi, 6500 chevron usually strongly marked with black and feet, 3 Sep 1940 (Roberts)—17M 18F (ANSP). yellow in males, may be similar in females or — km 60-75 E of San Luis Potosi, 5-6000 feet, mostly green and yellow. Lower marginal area with 13M 12F (Roberts) (ANSP). — Catorce, 28 Aug a row of dark spots. Uppper marginal area with 1935 (Roberts)—1M 2F (ANSP). — 8 miles E of two dark bands. Femur with a dark band anterior to San Luis Potosi (mail plaza) 6150 feet, 26 Aug 1959 narrowest part. Knee black in upper half, ivory in (Cohn and Cantrall #34)—2M 4F (Specimen H23) lower half. Lower edge of femur orange or red. (UMMZ). — 15 miles SW of Villa Hidalgo, 6100 Body measurements (mm).— Body length 15-19 feet, 20 Aug 1959 (Cohn and Cantrall #1)—4M 1F DANIEL OTTE 217
(Specimen H20) (UMMZ). 12. Philocleon ottei Fontana & Buzzetti 2007 Figures 4H, 12D, 14F, G, H, 16E, 17 11. Philocleon luceroae Fontana and Buzzetti 2007 Figures 4A, B, 12A-C, 14A-C, 16C, 17 Philocleon ottei Fontana & Buzzetti. 2007. Atti Acc. Rov. Agiati 8 7(B):113. Holotype male, Philocleon luceroae Fontana & Buzzetti. 2007. Mexico, Mexico Northeast, San Luis Potosí, Atti Acc. Rov. Agiati 8 7(B):108. Holotype betw. S. Martín de Abajo and S. Nicolás male. Mexico, Mexico Northeast, Querétaro, Tolentino. Paola Fontana Collection. Parque Nacional El Cimatario. Paulo Fontana Collection. Body measurements (mm).— Body length 17-19 (males), 22-27 (females). Hind femur length 10-11 Recognition.— Head: Antennae orange. Face (males), 12.5-15 (females). green or pale brown. Lateral lobe with a distinct Recognition.— Head: Antennae orange. Face black band along the lower margin (thus differing green or pale brown. Lateral lobe with upper dark from other members of the group). Lower pale band extending from front to back. Without a dark band on pronotum with two yellow anterior patches band along the lower margin (unlike luceroae). and one ivory posterior patch in male; with one Front and middle femora orange brown. Hind femur yellow anterior patch and two ivory patches in with very distinct black and yellow chevron. Lower female. Dorsum of abdomen greenish. Front and marginal area with a row of dark spots and below middle femora pink. Hind femur red or pink on that a band of red. upper marginal area. Medial area chevron green Specimens examined.— MEXICO: SAN LUIS and yellow. Femur without a dark area just anterior POTOSI: 10 miles NW of Rio Verde, 3550 feet, to the narrowest portion. Upper half of knee black, 25 Aug 1959 (Cohn and Cantrall #30)—2M 2F lower half greenish or grey. Hind tibiae blue green. (Specimen PH-26) (UMMZ). — 4.3 road miles Body measurements (mm).— Body length 17-19 W of Santa Catarina, 42 road miles WNW of Rio (males) 22-24 (females). Hind femur length 10-11 Verde, 4300 feet, 24 Aug 1959 (Cantrall and Cohn (males) 12-13 (females). #26)—1M (Specimen PH27) (UMMZ). — 19.5 Specimens examined.— MEXICO: JALISCO: miles E Rio Verde (on Hwy 70) 20 Nov 1970 NE, 10.5 miles NW of Leon (center of town) (Cohn and Cohn #101)—3M 2F (Specimen H40) (ridge summit) 11 Nov 1970 (Cohn and Cohn (UMMZ). — 6 miles W of San Jose de Gallinas #87)—6M 3F (Specimens H4, PH-22) (UMMZ). (22 miles WNW of Rio Verde) 3800 feet, 25 Aug — QUERETARO: Cuesta China, 4.2 miles E of 1959 (Cantrall and Cohn #27)—1M (Specimen Queretaro (south plaza), on old Hwy 45, 12 Nov H22) (UMMZ) (UMMZ). — 3.5 miles E of San 1970 (Cohn and Cohn #88)—(Specimens H5, Jose de Galinas (13 miles WNW of Rio Verde) 3450 PH-23) (UMMZ). — 9.4 miles N of Queretaro, feet, 25 Aug 1959 (Cantrall and Cohn #28)—2M 25 Aug 1965 (TJ Cohn #69)—(Specimen H44) 2F) (UMMZ). — km 160 E of San Luis Potosi, (UMMZ).— HIDALGO: 13 miles N of Atotonilco, 4000 feet, 8 Sep 1940 (Roberts)—M 7F (ANSP). 8-9000 feet, 3 Nov 1951, rocky, shrubby rim, desert — GUANAJUATO: 11 road miles E of San Luis de valley (JB Mosilmann #34)—2M (Specimens H1, La Paz (6900 feet), 29 Aug 1959 (Cohn and Cohn PH19) (UMMZ). — QUERETARO: 9.5 miles N #46)—2M (Specimen PH-33) (UMMZ). of Queretaro, 25 Aug 1965 (TJ Cohn #70)—2M (Specimen D24) (UMMZ). — GUANAJUATO: 11 road miles E of San Luis de La Paz (6900 feet), 29 Aug 1959 (Cohn and Cohn #46)—2M (Specimen PH-33) (UMMZ). — 12 road miles E of San Luis de La Paz (6750 feet), 28 Aug 1959 (Cohn and Cohn #38)—3M 1F (UMMZ). 218 REVISION OF PHILOCLEON
14. Philocleon erissa new species 13. Philocleon iropon new species urn:lsid:zoobank.org:act:00B1DD35-D51E-4F7C-A7A3- urn:lsid:zoobank.org:act:5AB9B204-E1C3-48D8-B51A- 2C101B6C2AF0 8802C328C4CC Figures 4F, 13D, 16A, 17 Figures 4J, K, 12D, 14I, 16D, 17 Holotype male.— Mexico, Coahuila, 25 miles Holotype male.— Mexico, Coahuila, 20 road S of Castaños, 3 Aug 1959, 3150 feet (TJ Cohn miles SE of Arteaga, 10 Aug 1959, 6400 feet #136)— Specimen H28. University of Michigan (TJ Cohn #162)—Specimen H24. University of Museum of Zoology. Michigan Museum of Zoology. Recognition.— Head: Antennae orange. Face Recognition.— Differs from the very similar yellowish. Lateral lobes with greenish bands. P. scudderi chiefly in the configuration of the male Mezozona with large dorsal and ventral ivory genitalia. Head: Antennae orange. Face yellowish. patches on lateral face. Upper ivory patch separated Lateral lobes streaked with black and yellow, upper from dorsal red area with a black band. Front and dark band interrupted. Front and middle femora middle femora usually orange. Hind legs: medial usually orange. Hind legs: medial area chevron area chevron usually strongly marked with black usually strongly marked with black and yellow in and yellow in males, may be similar in females or males, may be similar in females or mostly green mostly green and yellow. Lower marginal area with and yellow. Lower marginal area with a row of dark a row of dark spots. Uppper marginal area with spots. Uppper marginal area with two dark bands. two dark bands. Femur with a dark band anterior to Femur with a dark band anterior to narrowest part. narrowest part. Knee black in upper half, ivory in Knee black in upper half, ivory in lower half. Lower lower half. Lower edge of femur orange or red. edge of femur orange or red. Body measurements (mm).— Body length 17- Body measurements (mm).— Body length 15-19 19 (males), 21-24 (females). Hind femur length (males), 20-24 (females). Hind femur length 9-11 (males), (females). (males), 10.5-12 (females). Specimens examined.— MEXICO: Specimens examined.— MEXICO: COAHUILA: 25 miles S of Castaños, 3 Aug 1959, COAHUILA: 9 miles N of Concepcion del Oro, 3150 feet (TJ Cohn #136)—1M 1F (Specimen H28) 6 Aug 1959, 4950 feet (TJ Cohn # 147)—1M (UMMZ). — 26 miles S of Castaños plaza, 11 road (Specimen PH-2) (UMMZ). — 9 miles W of miles N of San Lazaro, 3040 feet, 17 Oct 1974 Saltillo, 18 miles E of La Rosa, 5500 feet, 27 Aug (Cohn and Cohn #84)—3M 1F (UMMZ). 1961 (Cantrall and Cohn #29)—1M (UMMZ). — 18.6 road miles S of Arteaga, 6850 feet, 20 August 1961 (Cantrall and Cohn #21)—4M 1F (Specimen PH-6) (UMMZ). — 20 road miles SE of Arteaga, 10 Aug 1959, 6400 feet (TJ Cohn #162)—6M 2F (Specimen H24) (UMMZ). — 40 miles SW of Saltillo, 5 Aug 1959, 5700 feet 9 (TJ Cohn #143)— 4M (Specimens H25, PH-1) (UMMZ). — 39 miles S of Castaños Plaza, 1.7 miles S of San Lazaro, 3600 feet, 17 Oct 1974 (Cohn and Cohn #83)—9M 11F (Specimen PH-34) (UMMZ). — 35 miles S of Castaños, 25 Sep 1958, 4000 feet (TJ Cohn #139 (Specimen PH-29) (UMMZ). — 20 miles W of Saltillo, 4300 feet, 16 Jul 1936 (Roberts)—32M 11F (ANSP). — NUEVO LEON: 6.6 miles S of Puerto Flores, 16.8 miles S of Arteaga, 6740 feet, 21 Aug 1961 (Cantrall and Cohn #26)—1M 1F (Specimen PH-43) (UMMZ). DANIEL OTTE 219
LITERATURE CITED
Eades, D. C., D. Otte, M. M. Cigliano & H. Braun. Orthoptera Species File Online. Version 2.0/4.0. [retrieval date].
Figure 1. Portraits of males belonging to the four Philocleon species groups. A, P. anomalus of the Anomalus Group. B, P, azumai of the Illa Group. C, P. nigrovittatus of the Nigrovittatus Group. D, P. scudderi of the Scudderi Group. DANIEL OTTE 221
Figure 2. Head and thorax and hind femur. A, B, Anomalus Group (A, anomalus; B, zygon). C-G, Illa Group (C, zima; D, cledon; E, azumai; F, axiton; G, illa). 222 REVISION OF PHILOCLEON
Figure 3. Head and thorax and hind femur of the Nigrovittatus Group. A-E, nigrovittatus; F-H, spatulatus. DANIEL OTTE 223
Figure 4. Head and thorax and hind femur of the Scudderi Group. A-B, luceroae; C, D, E, G, scudderi; F, erissa; H, ottei; J, K, iropon. 224 REVISION OF PHILOCLEON
Figure 5. Male aedeagus of the Anomalus Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed.A-C, anomalus; D, zygon. DANIEL OTTE 225
Figure 6. Epiphallus showing dorsal and posterior views. A-E, anomalus; F, G, zygon. 226 REVISION OF PHILOCLEON
Figure 7. Male aedeagus of the Illa Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed.A, illa; B, zima; C, cledon; D, axiton; E, azumai. DANIEL OTTE 227
Figure 8. Epiphallus showing dorsal and posterior views. A, illa; B, zima; C, cledon; D, E, axiton; F, G, azumai. 228 REVISION OF PHILOCLEON
Figure 9. Male aedeagus of the of P. nigrovittatus of the Nigrovittatus Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed (see Specimens section for localities). DANIEL OTTE 229
Figure 10. Male aedeagus of P. spatulatus of the Nigrovittatus Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed (see Specimens section for localities). 230 REVISION OF PHILOCLEON
Figure 11. Epiphallus showing dorsal and posterior views in the Nigrovittatus Group. A-F, nigrovittatus; G-M, spatulatus. DANIEL OTTE 231
Figure 12. Male aedeagus of the Scudderi Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed. A-C, luceroae; D, ottei; E, iropon. 232 REVISION OF PHILOCLEON
Figure 13. Male aedeagus of the Scudderi Group showing lateral, dorsal, and ventral views; right two figures showing aedeagus with the cingulum removed. A-C, scudderi; D, erissa. DANIEL OTTE 233
Figure 14. Epiphallus showing dorsal and posterior views in the Scudderi Group. A-C, luceroae; D, iropon; E, erissa; F, G, H, ottei; I, J, K, scudderi. 234 REVISION OF PHILOCLEON
Figure 15. Distribution maps of Philocleon species. A, anomalus; B, zygon; C, illa; D, zima; E, cledon; F, axiton; G, nigrovittatus; H, spatulatus; I, azumai. DANIEL OTTE 235
Figure 16. Distribution maps of Philocleon species. A, erissa; B, iropon; C, luceroae; D, scudderi; E, ottei. 236 REVISION OF PHILOCLEON
Figure 17. Cerci and furculae in all species.