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Introduction
The cockroach family Corydiidae Saussure, 1864 (Insecta: Blattodea) is characterized by the presence of an inflated clypeus, which is transversely divided into an anteclypeus and a postclypeus (Fig. 1); the wings, when present, have an anal area with a single fold; finally, there is the presence of two tubercles in the sclerite R2 of the male genitalia (Grandcolas, 1999; Estrada-Álvarez & Guadarrama, 2013). This family has a worldwide distribution and comprises around 260 species grouped in 44 genera (Beccaloni, 2014; Djernaes, 2018). Particularly, North America is home of 65 species which are grouped into eight genera and three subfamilies (Atkinson et al., 1991; Beccaloni, 2014; Estrada-Álvarez, 2013) (Table 1), none of which are of medical or economic significance.
Table 1 Checklist of species of the family Corydiidae recorded for North America, including its status (native, endemic or introduced) and state records for both Mexico and the USA, according mainly to Atkinson et al. (1991), Estrada-Álvarez (2013) and Hopkins (2014). Dubious records are marked with [?].
| Family CORYDIIDAE Saussure, 1864 |
| Subfamily CORYDIINAE Saussure, 1864 |
| Genus Arenivaga Rehn, 1903 |
| 1. Arenivaga adamsiHopkins, 2014. Mexico (Sinaloa, Sonora); USA (Arizona) |
| 2. Arenivaga akanthikosHopkins, 2014. Mexico (Sonora) |
| 3. Arenivaga alichenasHopkins, 2014. Mexico (Baja California) |
| 4. Arenivaga apacha (Saussure, 1893). USA (Arizona, New Mexico) |
| 5. Arenivaga apaeninsulaHopkins, 2014. Mexico (Baja California Sur) |
| 6. Arenivaga aquilaHopkins, 2014. Mexico (Guerrero, Morelos) |
| 7. Arenivaga belliHopkins, 2014. USA (Arizona, California, Nevada, Utah) |
| 8. Arenivaga bolliana (Saussure, 1893). Mexico (Durango, Nuevo Leon, Tamaulipas, Sonora [?]); USA (Florida [?], Texas) |
| 9. Arenivaga darwiniHopkins, 2014. Mexico (Sonora); USA (Arizona, California) |
| 10. Arenivaga delicataHopkins, 2014. USA (Arizona, California, Nevada, Utah) |
| 11. Arenivaga diaphanaHopkins, 2014. Mexico (Baja California) |
| 12. Arenivaga dnopherosHopkins, 2014. Mexico (Puebla) |
| 13. Arenivaga erratica Caudell, 1903. Mexico (Chihuahua, Coahuila, Durango); USA (Arizona, Nevada, New Mexico, Texas, Utah) |
| 14. Arenivaga estelleaeHopkins, 2014. Mexico (Baja California); USA (California) |
| 15. Arenivaga floridensis Caudell, 1918. USA (Florida) |
| 16. Arenivaga florilegaHopkins, 2014. Mexico (Hidalgo) |
| 17. Arenivaga gaiophanesHopkins, 2014. USA (California) |
| 18. Arenivaga galeanaHopkins, 2014. Mexico (Nuevo Leon) |
| 19. Arenivaga genitalis Caudell, 1918. Mexico (Sonora); USA (Arizona) |
| 20. Arenivaga grandiscanyonensisHopkins, 2014. USA (Arizona) |
| 21. Arenivaga grata Hebard, 1920. Mexico (Aguascalientes, Chihuahua, Coahuila, Durango, San Luis Potosi, Sonora); USA (Arizona) |
| 22. Arenivaga gumperzaeHopkins, 2014. Mexico (Coahuila, Durango, Nuevo Leon); USA (Texas) |
| 23. Arenivaga gurneyiHopkins, 2014. Mexico (Michoacan) |
| 24. Arenivaga haringtoniHopkins, 2014. USA (Arizona, Nevada) |
| 25. Arenivaga hebardiHopkins, 2014. Mexico (Sonora) |
| 26. Arenivaga hopkinsorumHopkins, 2014. Mexico (Sonora); USA (California, Arizona) |
| 27. Arenivaga hypogaiosHopkins, 2014. Mexico (Coahuila, San Luis Potosi); USA (Texas) |
| 28. Arenivaga impensaHopkins, 2014. USA (Arizona) |
| 29. Arenivaga investigata Friauf & Edney, 1969. Mexico (Baja California, Sonora); USA (Arizona, California, Nevada) |
| 30. Arenivaga mckittrickaeHopkins, 2014. USA (California) |
| 31. Arenivaga milleriHopkins, 2014. USA (California, Nevada) |
| 32. Arenivaga moctezumaHopkins, 2014. Mexico (Sonora) |
| 33. Arenivaga mortisvallisensisHopkins, 2014. USA (California) |
| 34. Arenivaga nalepaeHopkins, 2014. Mexico (Baja California); USA (Arizona, California, Nevada) |
| Genus Arenivaga Rehn, 1903 |
| 35. Arenivaga nickeliHopkins, 2014. Mexico (Baja California); USA (Arizona [?]) |
| 36. Arenivaga nocturnaHopkins, 2014. Mexico (Baja California, Coahuila) |
| 37. Arenivaga paganaHopkins, 2014. USA (Arizona, Nevada, Utah) |
| 38. Arenivaga paradoxaHopkins, 2014. Mexico (Baja California) |
| 39. Arenivaga pratchettiHopkins, 2014. USA (Arizona, California) |
| 40. Arenivaga pumilaHopkins, 2014. Mexico (Baja California Sur) |
| 41. Arenivaga rehniHebard, 1917. Mexico (Baja California, Baja California Sur); USA (California) |
| 42. Arenivaga riceiHopkins, 2014. USA (Texas) |
| 43. Arenivaga rothiHopkins, 2014. Mexico (Coahuila); USA (Texas) |
| 44. Arenivaga sequoiaHopkins, 2014. USA (California) |
| 45. Arenivaga tenaxHopkins, 2014. Mexico (Chihuahua); USA (Arizona, New Mexico, Texas) |
| 46. Arenivaga tonkawa Hebard, 1920. Mexico (Baja California, Coahuila, Sonora, Nuevo Leon, Tamaulipas); USA (Arizona, California, Colorado, Nevada, Oklahoma, Texas, Utah) |
| 47. Arenivaga trypherosHopkins, 2014. Mexico (Baja California); USA (Arizona, California) |
| 48. Arenivaga umbratilisHopkins, 2014. USA (Arizona) |
| Genus Eremoblatta Rehn, 1903 |
| 49. Eremoblatta atticolaEstrada-Álvarez & Rojas 2020. Mexico (Queretaro) |
| 50. Eremoblatta hirsutaHebard, 1917. Mexico (Baja California Sur) |
| 51. Eremoblatta subdiaphana (Scudder, 1902). USA (California, New Mexico, Nevada, Texas) |
| Genus Homoeogamia Burmeister, 1838 |
| 52. Homoeogamia mexicana Burmeister, 1838. Mexico (Ciudad de Mexico, Durango, Estado de Mexico, Guerrero, Guanajuato, Jalisco, Michoacan, Morelos, Oaxaca, Puebla, Sinaloa, San Luis Potosi, Veracruz) |
| Subfamily EUTHYRRHAPHINAE Handlirsch, 1925 |
| Genus Holocompsa Burmeister, 1838 |
| 53. Holocompsa azteca Burmeister, 1838. Mexico (Oaxaca, Veracruz); introduced to the USA (Arizona) |
| 54. Holocompsa nitidula (Fabricius, 1781). Mexico (Veracruz); introduced to the USA (Florida) |
| 55. Holocompsa scotaea Hebard, 1922. Mexico (Sinaloa) |
| 56. Holocompsa tolteca (Saussure & Zehntner, 1894). Mexico (Veracruz) |
| 57. Holocompsa zapoteca Saussure & Zehntner, 1894. Mexico (Yucatan) |
| Subfamily LATINDIINAE Handlirsch, 1925 |
| Genus Compsodes Hebard, 1917 |
| 58. Compsodes cucullatus (Saussure & Zehntner, 1894). Introduced to the USA (Florida, Texas) |
| 59. Compsodes mexicanus (Saussure, 1868) comb. rest. Mexico (Baja California Sur, Queretaro, Estado de Mexico, Veracruz) |
| 60. Compsodes schwarzi (Caudell, 1903). Mexico (Baja California Sur, Sinaloa); USA (Arizona, Florida, Texas) |
| Genus Latindia Stål, 1860 |
| 61. Latindia dohrniana Saussure & Zehntner, 1894. Mexico (Veracruz) |
| Genus Myrmecoblatta Mann, 1914 |
| 62. Myrmecoblatta rehniMann, 1914. Mexico (Hidalgo) |
| 63. Myrmecoblatta wheeleriHebard, 1917. USA (Florida) |
| Genus Paralatindia Saussure, 1868 |
| 64. Paralatindia aztecaSaussure, 1868. Mexico (Veracruz) |
| 65. Paralatindia hebardi (Estrada-Álvarez & Guadarrama, 2013) comb. nov. Mexico (Ciudad de Mexico, Estado de Mexico) |
The subfamily Corydiinae is represented in North America by three genera: Homoeogamia Burmeister, 1838; ArenivagaRehn, 1903; and EremoblattaRehn, 1903. The males of these three genera are macropterous (wings which surpass abdominal apex). The females of Arenivaga and Eremoblatta are streamlined and apterous (wingless), with a convex thorax and a sharp pronotum, this is an adaptation for a completely fossorial lifestyle, as it helps them “swim” through loose substrate (Bell et al., 2007). Less adapted “swimmers”, but still completely fossorial, are the females of Homoeogamia, which have tegmina and wings, yet remain flightless. This subfamily has a Holarctic distribution.
The subfamily Latindiinae includes the smallest members of this family, some less than 5 mm in length. The males of MyrmecoblattaMann, 1914 and ParalatindiaSaussure, 1868 are brachypterous (reduced tegmina and wings) (Deyrup & Fisk, 1984; Estrada-Álvarez & Guadarrama, 2013; Mann, 1914; Estrada-Álvarez & Rojas, 2020). Males of CompsodesHebard, 1917 and Latindia Stål, 1860 are macropterous. The females of Myrmecoblatta, Paralatindia, and Compsodes, are apterous; while those of Latindia are macropterous (Estrada-Álvarez & Guadarrama, 2013; Gutiérrez, 2012).
Finally, the subfamily Holocompsinae is represented in the region by the genus Holocompsa Burmeister, 1838; in this genus, both sexes are macropterous and capable of flight, and do not show sexual dimorphism, it is easily separated from the rest by the morphology of their tegmina (Qiu et al., 2020).
Materials and methods
For the making of the key, the works of Choate (2009), Estrada-Álvarez and Guadarrama (2013), Gutiérrez (2012), Hebard (1917a), Helfer (1987), Rehn (1903, 1950), Saussure and Zehntner (1893-1899) and Triplehorn and Johnson (2004) were consulted. The images were obtained from various authors, borrowed from articles (Estrada-Álvarez & Rojas, 2020; Gutiérrez, 2012; Hebard, 1917b; Saussure, 1870), or taken with a SONY α-6000 camera with a SEL30M35 macro lens. All the images were edited using the software GIMP© (Version 2.10).
The type specimens described by Saussure with type locality in Mexico were examined, they are deposited in the collection of the Muséum d'histoire naturelle de Genève, Geneva, Switzerland (MHNG). Other museum specimens were examined: Colección Entomológica (Entomological Research), Metepec, Estado de Mexico, Mexico (CER); Colección Nacional de Insectos, Instituto de Biología, UNAM, Ciudad de Mexico, Mexico (CNIN); Laboratorio de Ecología y Sistemática de Microartrópodos, Facultad de Ciencias, UNAM, Ciudad de Mexico, Mexico (LESM); Instituto de Ecología, Xalapa, Veracruz, Mexico (IEXA); Zoological Collections in the Museum of Biology of Lund University, Lund, Sweden (MZLU).
The checklist of the North American species of the family Corydiidae (Table 1) includes state records for both Mexico and the USA, and are based mainly in the works of Atkinson et al. (1991), Estrada-Álvarez (2013) and Hopkins (2014); other mentions for species of this family in the region were also consulted (Choate, 2009; Deyrup & Fisk, 1984; Estrada-Álvarez & Guadarrama, 2012; 2013; Estrada-Álvarez & Rojas, 2020; Estrada-Álvarez & Sormani, 2021; Hebard, 1917a; 1917b; 1921; Hollier et al., 2020; Mann, 1914; Núñez-Bazán et al., 2021; Peck & Beninger, 1989; Rehn, 1902; 1903; Saussure, 1864; 1870; Saussure & Zehntner, 1893-1894).
Results and discussion
I. Identification key for the subfamilies and genera of Corydiidae of North America
The following key is aimed to help in the identification process of adult specimens of Corydiidae found in Mexico and the USA to the genus level, it will likely fail if juveniles, or female Latindiinae, are examined instead. This key contemplates the taxonomic changes made below.
1 Well-developed tegmina often bearing yellowish-orange spots or bands; the proximal half is sclerotized while the distal half is membranous (Fig. 2) .... HOLOCOMPSINAE: Holocompsa
- Tegmina variable in development, if well-developed, they are completely coriaceous and lack said spots or bands ........................................................................................................................................... 2
2 Large sized species, measuring more than 10 mm in length .................................. CORYDIINAE ….. 3
- Minute species, measuring less than 10 mm in length ............................................. LATINDIINAE ..... 5
3 Pronotum with impressions due to the absence of setae (Fig. 3); both sexes winged (Fig. 3); total length about 30 mm, or more ......................................................................................... Homoeogamia
- Pronotum without said impressions; males macropterous, females apterous; total length 11-20 mm ......................................................................................................................................................................... 4
4 Males with subgenital plate with a spinous process on one side (Fig. 4); females with supra-anal plate not visible in dorsal view .............................................................................................. Eremoblatta
- Males with subgenital plate without said process; females with supra-anal plate visible in dorsal habitus ............................................................................................................................................... Arenivaga
5 Males with diagonal channel in tegmina (Fig. 5) ................................................................................................6
- Males without diagonal channel in tegmina ...................................................................................................... 7
6 Males brachypterous (Fig. 6A) ......................................................................................................... Paralatindia
- Males macropterous (Fig. 6B) ............................................................................................................. Compsodes
7 Males brachypterous (Fig. 6C); body about as wide as long (Fig. 6C); associated with ant’s nests (myrmecophilous) .............................................................................................................. Myrmecoblatta
- Males macropterous (Fig. 6D); body clearly longer than wide (Fig. 6D); not myrmecophilous ................................................................................................................................................................... Latindia
Figure 3 Homoeogamia mexicana, male, holotype, edited to show only the pronotum and tegminas; provided by the Biological Museum, Entomology, Lund University (available at https://www.flickr.com/photos/127240649@N08/50378520141/).
Figure 4 Eremoblatta atticola male, ventral aspect of the subgenital plate; borrowed from Estrada-Álvarez & Rojas (2020).
Figure 5 Compsodes perezgelaberti male, tegmina; arrow points to the diagonal channel; borrowed from Gutiérrez (2012).
Figure 6 Four species of Latindiinae, dorsal aspect. A) Paralatindia hebardi comb. nov. male, photo by Omar Sánchez. B) Compsodes schwarzi male, photo by Salvador Vitanza. C) Myrmecoblatta wheeleri male, borrowed from Deyrup & Fisk (1984). D) Latindia castanea Brunner von Wattenwyl, 1893, unknown sex, holotype (BMNH), photo by the trustees of the Natural History Museum, London.
II. Comments and taxonomic changes
Revising the type material of Paralatindia aztecaSaussure, 1868 and Myrmecoblatta hebardiEstrada-Álvarez & Guadarrama, 2013, we conclude that the latter is erroneously assigned to the genus Myrmecoblatta, being more akin with Paralatindia, therefore, it is transferred to this genus, generating Paralatindia hebardi (Estrada-Álvarez & Guadarrama, 2013) comb. nov.
Revising the type material of Latindia mexicanaSaussure, 1868 and Latindia tolteca Saussure & Zehntner, 1893, the synonymy proposed by Hebard, 1917 is confirmed. Furthermore, it is concluded that all the specimens referred by Saussure and deposited in the MHNG are males (contrary to Saussure, 1868, 1870; Saussure & Zehntner, 1893); with the collection and revision of new material, including females, it is confirmed that this species belongs to Compsodes Hebard, 1917: males and females of small size, less than 10mm, with hirsute integument and sexual dimorphism. Males macropterous. Spination type C0 sensuRoth (2003). With conspicuous diagonal channel in tegminas. Pronotum with medial carina. Supra-anal lamina with membranal window and convergent lateral margins and medial emargination. Sub-genital lamina with the projected space between cerci (species self-morphing); genital phalomere on the left side (sclerite L3). Female apterous. Therefore, L. mexicana is retransferred to the genus Compsodes, according to Hebard (1917a), generating Compsodes mexicanus (Saussure, 1868) (♂︎♀︎) comb. rest.
Revising the juvenile holotype of Homoeogamia brasiliana (Saussure, 1864) (MHNG) (Fig. 7), and comparing it with juveniles of Homoeogamia mexicana Burmeister, 1838, it was evident that there is no relation between these two species. When comparing with other juveniles of other Corydiidae of the MHNG, we noted that they were related to Polyphaga Brullé, 1835, especially to Polyphaga aegyptiaca (Linnaeus, 1758); with which we conclude that Polyphaga (Homoeogamia) brasiliana Saussure, 1864 is a JUNIOR SYNONYM of P. aegyptiaca and that the record for Brazil is anthropogenic, and hereby omitted. The records of Homoeogamia sp. n. Bolivar, 1890:137 (Juv.) [Caracas] and Homoeogamia sp. Rehn, 1903: 286 (Juv) [Buenos Ayres, Argentina] should be considered at the level of family: Corydiidae, this same conclusion was reached by Dr. A. B. Gurney (unpublished data).
Figure 7 Holotype of Polyphaga (Homoeogamia) brasilianaSaussure, 1864 jun. syn. Polyphaga aegyptiaca (Linnaeus, 1758). A) Dorsal habitus and labels. B) Dorsal habitus. C) Frontal aspect of the head. D) Frontal leg. E) Supra-anal plate. F) Sub-genital plate.
III. Synopsis of the family Corydiidae in North America
A complete checklist of species found in the region is present in Table 1.
Order BLATTODEA Brunner von Wattenwyl, 1882
Suborder BLATTARIA Burmeister, 1829 (sensu Klass & Meier, 2006)
[Blattodea ex. Isoptera (Li, 2019)]
Superfamily CORYDIOIDEASaussure, 1864
Family CORYDIIDAESaussure, 1864
Subfamily CORYDIINAESaussure, 1864
I. Genus Arenivaga Rehn, 1903
Figure 8 Two species of Arenivaga. A) Arenivaga bolliana male, lectotype (MHNG), dorsal habitus, with wings spread. B) Arenivaga bolliana female, paralectotype (MHNG), dorsal habitus. C) Arenivaga apacha male, lectotype (MHNG), dorsal aspect of pronotum. D) Arenivaga apacha male, lectotype (MHNG), frontal aspect of head. E) Arenivaga apacha female, paralectotype (MNHG), frontal aspect of the head. F) Arenivaga apacha female, paralectotype (MNHG), supra-anal plate.
Homoeogamia (Arenivaga) Rehn, 1903b: 181, 185 [subgen. nov.].
Arenivaga Caudell, 1913: 605 [stat. nov. genus].
ArenivagaHopkins, 2014: 17 [revision].
Type species: Homoeogamia (Arenivaga) bollianaSaussure, 1893 [=Arenivaga bolliana], by original designation Rehn (1903c: 185)
Brief diagnosis: Corydiinae with sexual dimorphism: macropterous males and apterous females. Both sexes resemble Eremoblatta; males differ in lacking a spinous process in one lateral margin of the subgenital plate; females differ in having visible supra-anal plate in dorsal habitus.
Distribution and biogeography: The species of this genus are distributed in the USA and Mexico, continuously from the southwest of the USA to the northern region of Mexico, with presence in the south-central region of Mexico and in an isolated region of Florida, USA. Biogeographically, it is present in three regions of America, it is more predominant in the Nearctic region, but it is also found in the Sierra Madre Oriental and in the Neotropical region.
Bionomy: Fossorial females and nymphs; sandy habitats; feeding on mycorrhizal fungi, desert foliar debris and seeds collected by mammals, with which they often coexist (Hopkins, 2014). Females are more active in summer (Hopkins, 2014); they live primarily in xeric scrubland. Positive phototropism is reported only in males (Hopkins, 2014).
Species reported for North America: 48, see Table 1 (species 1-48).
Notes: Several unidentified species of this genus have been mentioned in the literature (Estrada-Álvarez, 2013), even after the keys of Hopkins (2014) were published (Cueto-Medina et al., 2015).
II. Genus Eremoblatta Rehn, 1903
Figure 9 Eremoblatta atticola, borrowed from Estrada-Rojas & Rojas (2020). A) Male holotype, outline of dorsal habitus. B) Female paratype, outline of dorsal habitus. C) Female paratype, supra-anal plate. D) Female paratype, subgenital plate.
Homoeogamia (Eremoblatta) Rehn, 1903b: 189 [subgen. nov.].
Eremoblatta Caudell, 1913: 605 [stat. nov. genus].
Type species: Homoeogamia (Eremoblatta) subdiaphana (Scudder, 1902) [=Homoeogamia subdiaphana Scudder, 1902]; by monotypy (Rehn, 1903: 189).
Brief diagnosis: Corydiinae with sexual dimorphism: macropterous males and apterous females. Both sexes resemble Arenivaga; males differ in having a spinous process in one lateral margin of the subgenital plate; females differ in having the supra-anal plate hidden in dorsal habitus.
Distribution and biogeography: The species are distributed in southwestern USA and Mexico. Biogeographically it is clearly a Nearctic genus.
Species reported for North America: Three, see Table 1 (species 49-51).
III. Genus Homoeogamia Burmeister, 1838
Figure 10 Homoeogamia mexicana A) Male holotype, edited to show the frontal aspect of the head, provided by the Biological Museum, Entomology, Lund University (available at https://www.flickr.com/photos/127240649@N08/50378699032/in/photostream/). B) Female from Estado de Mexico, Mexico, dorsal habitus. C) Female from Estado de Mexico, Mexico, outline of the frontal aspect of the head. D) ♀︎ from Estado de Mexico, Mexico, supra-anal plate. E) Female from Estado de Mexico, Mexico, subgenital plate.
Heterogamia (Homoeogamia) Burmeister, 1838: 490 [subgen. nov.].
Homoeogamia Brunner von Wattenwyl, 1865: 360 [status nov.].
Type species: Homoeogamia mexicana Burmeister, 1838 [=Heterogamia (Homoeogamia) mexicana], by monotypy (Burmeister, 1838: 490).
Brief diagnosis of the genus: Corydiinae with sexual dimorphism, both sexes winged, macropterous males and mesopterous (tegmina and wings reach abdominal apex) females. Females possess a divided subgenital plate.
Distribution and biogeography: Endemic of Mexico, present in 13-16 states (Estrada-Álvarez, 2013; Núñez-Bazán et al., 2021). Present in the Nearctic and Neotropical regions, as well as in the Mexican Transition Zone; most of the records of this species are within the Trans-Mexican Volcanic Belt; the records from the Nearctic region correspond to the lower part of the Mexican Altiplano; the Neotropical records are within the Gulf of Mexico, the Balsas Basin, and the Mexican Pacific Coast.
Bionomy: Epigean species, with cave-dwelling records; the males revised have a layer of soil on the pronotum and legs, suggesting a fossorial lifestyle. They are distributed from temperate forests to mesophyll forests, deciduous forests, and evergreen forests, as well as xerophilous scrubland; it is also present in several urban areas of the country, mostly in temperate climate, but also in hot and dry climates. All of the above shows the great adaptability of this species.
Species reported for North America: One, see Table 1 (species 52).
Subfamily HOLOCOMPSINAE Rehn, 1951
IV. Genus Holocompsa Burmeister, 1838
Figure 11 Two species of Holocompsa. A) Holocompsa zapoteca male, holotype (BMNH), dorsal habitus, photo by Edward Baker. B) Holocompsa tolteca female, holotype (BMNH), tegmina. C) Holocompsa tolteca female, holotype (BMNH), extended wing.
Corydia (Holocompsa) Burmeister, 1838: 491 [subgen. nov.].
HolocompsaSaussure, 1864c; 150 [status nov. genus].
Holocompsa Brunner von Wattenwyl, 1865: 346.
HolocompsaSaussure & Zehntner, 1893: 108, (104-105, in key).
Pseudoholocompsa (sic) [Pseudoholocompsa] Shiraki, 1931: 176 [gen. nov.] [jun. syn. Princis, 1963: 93].
Holocompsa Princis, 1963: 93 [syn. nov. Pseudoholocompsa].
Type species: Blatta nitidula Fabricius, 1781 [=Holocompsa nitidula], by designation by Burmeister, 1838: 491.
Brief diagnosis of the genus: Both sexes winged, not sexually dimorphic. Easily recognizable by the tegminas which have their proximal half sclerotized while the distal half is membranous.
Distribution and biogeography: Circumtropical. In America, it is present in the USA, Mexico, Guatemala, Panama, Costa Rica, Guyana, French Guyana, and Brazil. In Mexico the records come from the Neotropical region.
Bionomy: Cave-dwelling and epigean records.
Species reported for North America: Five, see Table 1 (species 53-57).
Subfamily LATINDIINAE Handlirsch, 1925
V. Genus Compsodes Hebard, 1917
Figure 12 Compsodes mexicanus. A) Male lectotype (MNHG), dorsal habitus. B) Male lectotype (MNHG), ventral habitus. C) Male lectotype (MNHG), supra-anal plate. D) Female from Estado de Mexico, Mexico, subgenital plate. E) Female from Estado de Mexico, Mexico, dorsal habitus.
CompsodesHebard, 1917a: 208 [gen. nov.].
CompsodesGutiérrez, 2012: 54.
Type species: Latindia delicatulus Saussure & Zehntner [=Compsodes delicatulus], by original designation (Hebard, 1917a: 209).
Brief diagnosis of genus: Latindiinae with sexual dimorphism, macropterous males and apterous females; arolium vestigial.
Distribution and biogeography: Discontinuous distribution, probably due to lack of collections or due to the size of the species of this genus. Biogeographically they are found in both the Nearctic and the Neotropical region.
Bionomy: Compsodes mexicanus (Saussure, 1868) comb. rest. is found in rotten logs.
Species reported for North America: Three, see Table 1 (species 58-60).
VI. Genus Latindia Stål, 1858
Latindia Stål, 1858: 311 [gen. nov.]
Zatindia (sic) Finot, 1897: 211.
Type species: Latindia maurella Stål, 1860, by monotypy.
Brief diagnosis of genus: Latindiinae without sexual dimorphism; males without hyaline window; tarsal claws simple; arolium present or absent.
Distribution and biogeography: Mexico to South America.
Species reported for North America: One, see Table 1 (species 61).
VII. Genus Myrmecoblatta Mann, 1914
MyrmecoblattaMann, 1914: 172 [gen. nov.].
Type species: Myrmecoblatta rehniMann, 1914; by monotypy.
Brief diagnosis of the genus: Myrmecophilous (associated with ant’s nests) Latindiinae with robust bodies. Sexually dimorphic, males brachypterous, females apterous. Tarsal claws simple and symmetrical, pulvillus and arolium absent. Females with frontal “Y” shaped suture (Mann, 1914; Fisk et al., 1976).
Distribution and biogeography: USA, Mexico, Guatemala, Nicaragua and Costa Rica.
Bionomy: Associated with ant’s nests (see Estrada-Álvarez & Rojas, 2020).
Species reported for North America: Two, see Table 1 (species 62-63).
Genus VIII. Paralatindia Saussure, 1868
Figure 13 Paralatindia azteca. A) Male lectotype (MHNG), dorsal habitus. B) Male lectotype (MHNG), ventral habitus. C) Male lectotype (MHNG), supra-anal plate. D) Female paralectotype (MHNG), subgenital plate. E) Female paralectotype (MHNG), dorsal habitus.
ParalatindiaSaussure, 1868: 100 [gen. nov].
Latindia (Paralatindia) Saussure, 1870: 112 [status nov. subgenus].
Paralatindia Saussure & Zehntner, 1894: 113.
Type species: Paralatindia aztecaSaussure, 1868, by monotypy.
Brief diagnosis of the genus: Latindiinae with sexual dimorphism, brachypterous males and apterous females.
Distribution and biogeography: Endemic to Mexico, records in the Trans-Mexican Volcanic belt.
Bionomy: Paralatindia hebardi (Estrada-Álvarez & Guadarrama, 2013) comb. nov. were collected in humid places.
Species reported for North America: Two, see Table 1 (species 64-65).
MATERIAL EXAMINED: Arenivaga bolliana Saussure, 1893 [=Homoeogamia (Arenivaga) bolliana], 1♂ lectotype, 1♀, 1 juv, paralectotype, Dallas, Texas, USA (MHNG). Arenivaga apacha (Saussure, 1893) [=Homoeogamia apacha], 1♂ lectotype, 1♂, 1♀ paralectotype, Chihuahua, Mexico, Mr. Corcelle coll. (MHNG). Arenivaga aquila Hopkins, 2014 1♀ allotype (with ootheca), 2♀♀, 1♀ juv. REBIOSH, El Limon de Cuauchichinola, Morelos, Mexico, pitfall trap, X/2020, Reinier Núñez coll. (CER). n. rec.; 1♀, 3.8 km west of Santa Cruz, Mexico, bait trap, human feces, 27/X/2005, Vaz de Melo, F. coll. (IEXA) n. rec.; 1♂, Km. 10 Cacahuamilpa-Taxco road, Acuitlapan, Guerrero, Mexico 12/VI/2001, H. Brailovsky and E. Barrera cols. (CNIN) n. rec.; 2♂♂, km. 24 Taxco-Iguala road, 3 km. Poniente de Mexcaltepec, Guerrero, Mexico, H. Brailovsky and E. Barrera colls. (CNIN). Arenivaga dnopheros Hopkins, 2014 1♀, 5 km north of Totoltepec, Puebla, Mexico, bait trap with human feces, 20/XII/2005, Vaz de Melo, F. coll. (IEXA). Arenivaga sp. (pos. Arenivaga florilega Hopkins, 2014) [det. Eremoblatta sp. n. hirsuta Hebard; det. F. W. Fisk 1983; ID err]; 2♀♀, km. 75.5 in road Toliman-Higuerillas, Queretaro, Mexico (III-C), in detritus of Atta mexicana, Larrea scrubland, 29/VIII/1982, P. Rojas coll. (CBS). Compsodes delicatulus (Saussure & Zehntner, 1894) 1♂ holotype, Zapote, Guatemala, G. C. Champion coll. (BMNH-876601), photos revised. Compsodes mexicanus Saussure, 1868 [=Latindia mexicanaSaussure, 1868], 1♂ lectotype (designated here) (MHNG), revised; 2♂♂ paralectypes, Moyoapam (sic), Mexique, Sumichrast coll. (MHNG), revised; 2♂♂ (1♀ (sic)) Sintypes. Moyoapam (sic), Mexique, Sumichrast coll. (MHNG). Latindia tolteca Saussure & Zehntner, 1893, revised; 1♂ (♀ (sic)) holotype, Moyoap, Mexique, Sumichrast coll. (MHNG), revised; 1♂, Tuxpan, Mich., 19/VII/1962, (CNIN); 4♂♂, Alpizahua, Jalisco, Mexico 14/IX/1950; (CNIN); 1♂, San Cristobal de las Casas, Chiapas, Mexico, 2100 m, 20/XII/1955, P. B. Rotger coll. (ex. C. Bolívar) (CNIN); 3♂♂, 6♀♀, 2 juv, Metepec, Cerro de los Magueyes, Estado de Mexico, Mexico, 18/IX/2018, J. C. Estrada-Álvarez & Balam Estrada F. colls. (CER); 1♂, rural road in Xalapa Enríquez and El Castillo near La Laguna de Casa Blanca. Cafetal-Cañaveral, Veracruz, Mexico, in the nest of Camponotus, under a stone in company of big gastropods, manual collecting, 2002, H. C. G. Sormani coll. (IEXA); 1♂ Xalapa Enríquez, Infonavit Sumidero, Zona Urbana, in the wall of a garden, manual collecting, I/2008, H. C. G. Sormani coll. (IEXA); 1♂, deviation Higuerillas road to San Juan del Rio-Jalpan, Queretaro, Mexico. (VIII-B), in detritus of Atta mexicana, Larrea scrubland, 31/V/1983, P. Rojas coll. (CBS) [det. F. W. Fisk, 1983]; 2♂♂, Mexico, deviation Higuerillas road to San Juan del Rio-Jalpan, Queretaro, Mexico. (VIII-A), in detritus of Atta mexicana, Larrea scrubland, 11/IV/1982, P. Rojas coll. (CBS) [det. F.W. Fisk, 1983]. Compsodes schwarzi (Caudell, 1903) 6♂♂, Estacion de Biología Chamela, Jalisco, Mexico Cuenca 1, Malaise trap; (2♂♂ 7-12/X/1992; 2♂♂ 18-23/VIII/1992; 1♂ 21-26/X/1992; 1♂ 28/XI-03/XII/1991), A. Rodríguez coll. (LESM); 2♂♂ Est. Biol. Chamela?, Jalisco, Mexico, W-4A, fumigation (1♂ 14/V/1993; 1♂ 25/XI/1993), (LESM); 1 juv. Estacion de Biología Chamela, Jalisco, Cuenca 4A, funnel 8, fumigation, 18/VIII/1992, A. Pescador, A. Rodríguez, T. A. Gómez, Chen Yan cols. (LESM); 4♂♂, Alpizahua, Jalisco, 14/IX/1950 (CNIN). Eremoblatta atticola Estrada-Ávarez & Rojas, 2020 1♂ holotype, 1♂ and 1♀ paratype, km. 75.5 Toliman-Higuerillas, Queretaro, Mexico, 29/VIII/1982, P. Rojas coll. (IEXA); 1♂ 2♀, paratypes, km. 75.5 Toliman-Higuerillas, Queretaro, Mexico, 27/I/1980, P. Rojas coll. (CER); 1♂ paratype, Cadereyta, Queretaro, Mexico, 14/XI/1991, H. Braylovsky and E. Barrera colls. (CNIN). Eremoblatta hirsuta Hebard, 1917 1♂ holotype, Sierra El Tasti (Tosti (sic) en Hebard, 1917) [=Cerro El Taste], Baja California, Mexico: oct. 93; Gust. Eisen coll. (ANSP) fototypes Heidi Hopkins, in Beccaloni (2014). Eremoblatta subdiaphana (Scudder, 1902) 1♂ topotype, Heidi Hopkins, in Beccaloni (2014). Holocompsa azteca (Saussure, 1862) 1♂ holotype, Moyoapan, Veracruz, Mexico; 2♂♂ 6♀♀ Mexico and Costa Rica (MHNG), revised; 2♀♀ Xalapa Enriquez, Coapexpan, induced grassland, in the hole in the base of a tree, manual collecting, H. C. G. Sormani & V. J. A. Ángeles colls. (IEXA); 1♀ Orizaba, Veracruz, Mexico, No. 19 (CNIN); 1♀ km. 84 Carr. Valladolid Rio, Yucatan, Mexico, 30/IV/1982, A. Barrera coll. (CNIN). Holocompsa tolteca (Saussure & Zehntner, 1894) Plan del Rio, Emiliano Zapata, Veracruz, Coconut plantation, under the dead log of a coconut tree, manual collecting, III/2003 H. C. G. Sormani coll. (IEXA); Xalapa Enriquez, Coapexpan, Veracruz induced tree, at the hollow base of a Liquidambar, manual collecting, V/2004, H. C. G. Sormani, V. J. A. Ángeles colls. (IEXA). Holocompsa zapoteca Saussure & Zehntner, 1894 1♂ Tekax, Cueva Actun Chocantes, Yucatan, Mexico, 19/IX/1995, O. García coll. (LESM). Homoeogamia azteca Saussure, 1893 2♂♂, 3♀♀, 4 juv types, Omilteme, Guerrero, Mexico, VII, H. H. Smith coll. (MHNG), revised. Homoeogamia mexicana Burmeister, 1838 1♂ type sensu Princis 1954. Mexico, Doppe(?) (MZLU), photos revised; 1♂ San Vicente, Guerrero, Mexico [10-960-2 ?], Ex. Colección Bolívar (CNIN); 1♂ Ciudad Universitaria, Ciudad de Mexico, Mexico, 14/XI/2017, G. Salgado coll. (CNIN); 1♂ Santiago de Anaya, Hidalgo, Mexico, Sección Norte, 24/VI/2006, C. Mayorga coll. (CNIN); 3♂♂ Instituto de Biología, UNAM, Ciudad de Mexico, 16/V/2006, C. Mayorga coll. (CNIN); 1♂ Instituto de Biología, UNAM, Ciudad de Mexico, 09/VI/2004, E. Mejorada coll. (CNIN); 1♂ Ciudad Universitaria, Ciudad de Mexico, Mexico, 14/VIII/2001, C. Mayorga coll. (CNIN); 1♂ Uruapan, Michoacan, Mexico (CNIN); 1♂ no data (CNIN); 4♂♂ Coyoacac, Ciudad de Mexico, 05/VI/1940 (CNIN); 1♂ Jardin Botánico, Ciudad Universitaria, Ciudad de Mexico, Mexico, 07/VI/1984, L. Vázquez coll. (CNIN); 1♂ Tacubaya, Ciudad de Mexico, Mexico (CNIN); 1♂ Uruapan, Michoacan, Mexico VI/1942 (CNIN); 1♂ Santiago de Anaya, Centro, Seccion Norte, Hidalgo, Mexico (20° 23´ 04´´ N, 98° 57´ 53´´ O); 04/X/2014; C. Mayorga coll. (CNIN); 1♂ Sierra de Guadalupe, Estado de Mexico, Mexico, VIII-XII/2017 (CNIN); 1♂ No data (LESM); 1♀ Metepec, Estado de Mexico, Mexico, 12/VII/2017, J. C. Estrada-Álvarez coll. (CER); 1♀ km. 18 road Donguiño, Hidalgo, Mexico, 04/VI/1999, E. Barrera & H. Braylovsky coll. (CNIN); 1♀ México, 05/IX/2000, A. Castillo coll. (CNIN); 1♀ Actopan, Hidalgo, Mexico, 1936l, C. C. Hoffmann coll. (CNIN); 1♀ Zempoala, Hidalgo, Mexico.; VII/1940, D. Zavaleta coll. (CNIN); 1♀ Alambaro (sic) [=Acámbaro], Guanajuato, Mexico (CNIN). 1♀ Uruapan, Michoacan, Mexico, VII/2000, Romano coll. (CNIN). Paralatindia azteca Saussure, 1868 1♂ lectotype (designed here), 6 ♂♂ 3 ♀♀ 4 juv. paralectotypes, Moyoapam (sic) [Moyo, Moyoap], Mexique, Sumichrast coll. (MHNG), revised; 2♂♂ El Volcancillo-Toxtlacoaya, pine forest, manual collecting, V/2004; H. C. G. Sormani & V. J. A. Ángeles, colls. (IEXA). Paralatindia hebardi (Estrada-Álvarez & Guadarrama, 2013) 1♂ Lerma, Estado de Mexico, Mexico 19/XI/2019, Eduardo Serafín coll. (CER); 1♂ Metepec, Barrio de San Mateo, in a house, 21/VII/2018, J. C. Estrada-Álvarez coll. (CER); 1♂ Toluca, Estado de Mexico, Mexico, 23/VII/2018, J. C. Estrada-Álvarez coll. (CER); 1♀ 1♂ juv. 1♀ juv. 5 juv and 1 ootheca, Colonia Morrelos 1ra. Sec. Toluca, Estado de Mexico, Mexico; 30 /IX/2013; Jorge Armando Mata González and Gretta P. Reyes Anzaldo colls. (CER); Polyphaga aequalis Walker, 1870 1♂, Beccaloni, 2019, BMNH, photos revised. Polyphaga (Homoeogamia) brasiliana Saussure, 1864 1 juv, holotype [nymph], Bresil, M. Sordet coll. (MHNG). Polyphaga aegyptiaca (Linnaeus, 1758) 16♂♂ 18♀♀ 29 juv (♂♂ and ♀♀) multiple locations (MHNG).
