Ripipterygidae

Ripipterygidae is a family of insects in the order Orthoptera. Members of the family are commonly known as mud crickets.

Mud crickets
Temporal range: Cenomanian-Recent
Ripipteryx mopana
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Caelifera
Infraorder: Tridactylidea
Superfamily: Tridactyloidea
Family: Ripipterygidae
Ander, 1939
Genera
  • †Archaicaripipteryx Xu, Zhang, Jarzembowski & Fang, 2020
  • Ripipteryginae
    • Mirhipipteryx Günther, 1969
    • Ripipteryx Newman, 1834
Synonyms

Rhipipterygidae Ander, 1939 (incorrect original spelling)

Description
Ripipteryx sp. (possibly R. biolleyi), Cartago, Costa Rica

Ripipterygids are small, often dark-colored, cricket-like orthopterans, between 3 and 14 mm in length. They closely resemble the related tridactylids. Like tridactylids, they have greatly expanded hind femora, and have the ability to swim and jump from the surface of water. They can be distinguished from tridactylids by their uninflated tibiae on the middle pair of legs, unsegmented cerci, rows of comblike teeth on the epiproct, and setae at the tips of the cerciform lobes on the paraproct, as well as through characters of the genitalia.[1]

Species in the genus Ripipteryx are typically black or dark brown and often metallic; many are boldly colored or strikingly patterned, with striking white, yellow, and/or red markings.[2] Members of the genus Mirhipipteryx are typically smaller and more drably colored.[1]

Distribution and habitat
Ripipteryx limbata, Cartago, Costa Rica

Ripipterygids are restricted to the Neotropics. They can be found from southern Mexico south to central South America. Like tridactylids, they are typically found in riparian areas, on bare soil, sand, and mud, as well as on rocks, and – in some species – low vegetation in, near, or above water.[3][4] In at least some species, individuals are not distributed evenly throughout appropriate habitat, but rather are found in groups.[4]

Biology

Ripipterygids have been comparatively little-studied, and many aspects of their biology are poorly known, with behavioral observations having only been made on a handful of species.[5]

Locomotion

Like tridactylids, ripipterygids walk quadrupedally using only their first two pairs of legs, and the hind legs are only used for jumping and swimming. Ripipterygids fly when disturbed, often landing on water and swimming back to shore.[4]

Diet and foraging

Ripipterygids are herbivorous, and have been recorded feeding on a variety of different plants, as well as foraging on the ground.[6][4][7] Foraging ripipterygids may leave networks of shallow tracks in sandy or muddy soil near waterways.[4]

Burrowing

At least some ripipterygids build short burrows or oval-shaped cells in clay soil or sand, on both flat ground and in vertical banks.[3][4][8] These burrows are used for temporary shelter, and may also be used for protection while molting.[4]

Daily activity pattern

Species vary in their activity patterns; some are most active during dawn and dusk, and others are active throughout the day even in direct sunlight.[3][8] One species studied was nocturnal.[3]

Breeding

In Ripipteryx notata in Uruguay (near the southern limit of the family's geographic distribution), females oviposit eggs singly in bare soil in spring and summer. Nymphs are present in summer and early autumn (whereas adults are present year-round).[6][4] Unlike in grasshoppers, embryonic molt has not been observed to occur in ripipterygids.[6]

Predation

Fish have been recorded preying on swimming ripipterygids.[4]

Taxonomy and evolution
Ripipteryx diegoi
Ripipteryx gorgonaensis
Ripipteryx guacharoensis

Ripipterygids, tridactylids, and sandgropers comprise the superfamily Tridactyloidea within the suborder Caelifera (grasshoppers). The tridactyloids are sister to the remainder of Caelifera. Within Tridactyloidea, ripipterygids are sister to tridactylids; the two families are estimated to have diverged between 150 and 175 million years ago.[9]

Ripipterygids were initially included in the family Tridactylidae. They were first elevated to family rank (as Rhipipterygidae) in 1939.[10] The classification of the family was extensively revised in a series of works by Kurt Günther between the 1960s and 1990s.[11][12][13][14][15][16][17][8] However, the taxonomic diversity of the family, particularly in Central America, is still incompletely known.[18][5]

Two extant genera of ripipterygids are recognized: Mirhipipteryx and Ripipteryx. Since the 1960s, the genus Ripipteryx has been divided into five species groups, based primarily on characters of the male genitalia. However, there is evidence that these groups are not monophyletic.[2][19][20] In addition, Ripipteryx itself may be paraphyletic with respect to Mirhipipteryx, suggesting additional taxonomic revision of the family is required.[19]

Both Mirhipipteryx and Ripipteryx have representatives distributed throughout the geographic distribution of the family, as do all of the Ripipteryx species groups apart from the Scrofulosa group, the members of which are all restricted to Central America.[3]

List of species

The following recent genera, species and subspecies of Ripipterygidae are recognized by OSF:[10]

  • Mirhipipteryx Günther, 1969
    • Mirhipipteryx acuminata Günther, 1969
    • Mirhipipteryx andensis Günther, 1969
    • Mirhipipteryx biloba (Hebard, 1928)
      • M. b. aberrans Günther, 1989
      • M. b. biloba (Hebard, 1928)
      • M. b. chiriquensis Günther, 1969
      • M. b. incurvata Günther, 1989
      • M. b. riofriensis Günther, 1969
      • M. b. sevillensis Günther, 1969
    • Mirhipipteryx columbiana (Günther, 1963)
      • M. c. columbiana (Günther, 1963)
      • M. c. tenaensis Günther, 1969
    • Mirhipipteryx disparilobata Günther, 1989
    • Mirhipipteryx hebardi (Chopard, 1931)
    • Mirhipipteryx hondurica Günther, 1969
    • Mirhipipteryx hubbelli Günther, 1969
    • Mirhipipteryx imperfecta Günther, 1989
    • Mirhipipteryx lilo Günther, 1969
      • M. lil. granchacensis Günther, 1977
      • M. lil. lilo Günther, 1969
    • Mirhipipteryx lineata Günther, 1989
      • M. lin. anchicayensis Günther, 1994
      • M. lin. lineata Günther, 1989
    • Mirhipipteryx lucieni Günther, 1969
    • Mirhipipteryx magdalenensis Günther, 1969
    • Mirhipipteryx panamica Günther, 1969
    • Mirhipipteryx peruviana (Saussure, 1896)
    • Mirhipipteryx phallica Günther, 1969
    • Mirhipipteryx pronotopunctata Günther, 1969
    • Mirhipipteryx pulicaria (Saussure, 1896)
      • M. pu. interposita Günther, 1969
      • M. pu. pulicaria (Saussure, 1896)
    • Mirhipipteryx schuchmanni Günther, 1994
    • Mirhipipteryx striatipes (Chopard, 1954)
    • Mirhipipteryx triangulata Günther, 1969
    • Mirhipipteryx unispinosa Günther, 1989
    • Mirhipipteryx variabilis Günther, 1969
    • Mirhipipteryx venezuelensis Günther, 1976
  • Ripipteryx Newman, 1834
    • Limbata-Marginata group
      • Ripipteryx boliviana Bruner, 1916
      • Ripipteryx cruciata Bruner, 1916
      • Ripipteryx cyanipennis Saussure, 1874
      • Ripipteryx furcata Günther, 1976
      • Ripipteryx hydrodroma Saussure, 1896
      • Ripipteryx lawrencei Günther, 1969
      • Ripipteryx limbata (Burmeister, 1838)
      • Ripipteryx marginata Newman, 1834
      • Ripipteryx notata (Burmeister, 1838)
      • Ripipteryx ornata Chopard, 1927
      • Ripipteryx rivularia Saussure, 1896
      • Ripipteryx saopauliensis Günther, 1969
      • Ripipteryx trilobata Saussure, 1874
      • Ripipteryx trimaculata Günther, 1969
    • Crassicornis group
      • Ripipteryx antennata Hebard, 1924
      • Ripipteryx atra Serville, 1838
      • Ripipteryx capotensis Günther, 1970
      • Ripipteryx crassicornis Günther, 1969
      • Ripipteryx gorgonaensis Baena-Bejarano & Heads, 2015
      • Ripipteryx laticornis Günther, 1963
      • Ripipteryx nodicornis Hebard, 1924
    • Forceps group
      • Ripipteryx carbonaria Saussure, 1896
      • Ripipteryx diegoi Baena-Bejarano, 2015
      • Ripipteryx ecuadoriensis Günther, 1962
      • Ripipteryx forceps Saussure, 1896
      • Ripipteryx nigra Günther, 1963
      • Ripipteryx paraprocessata Günther, 1989
      • Ripipteryx pasochoensis Heads, 2010
      • Ripipteryx processata Günther, 1969
    • Marginipennis group
      • Ripipteryx amazonica Günther, 1969
      • Ripipteryx brasiliensis Günther, 1969
      • Ripipteryx bruneri Chopard, 1920
      • Ripipteryx difformipes Chopard, 1956
      • Ripipteryx femorata Chopard, 1956
      • Ripipteryx guacharoensis Baena-Bejarano & Heads, 2015
      • Ripipteryx insignis Chopard, 1937
      • Ripipteryx luteicornis Chopard, 1920
      • Ripipteryx marginipennis Bruner, 1916
      • Ripipteryx sturmi Günther, 1963
      • Ripipteryx vicina Chopard, 1956
    • Scrofulosa group
      • Ripipteryx biolleyi Saussure, 1896
      • Ripipteryx mediolineata Günther, 1969
      • Ripipteryx mexicana Saussure, 1859
      • Ripipteryx mopana Heads & Taylor, 2012
      • Ripipteryx saltator Saussure, 1896
      • Ripipteryx saussurei Günther, 1969
      • Ripipteryx scrofulosa Saussure, 1896
      • Ripipteryx tricolor Saussure, 1896

In addition, two fossil species have been described, both based on individuals preserved in amber in regions of the world where the family no longer occurs: Mirhipipteryx antillarum Heads, 2010 (from early Miocene amber from Hispaniola[1]), and Archaicaripipteryx rotunda Xu, Zhang, Jarzembowski & Fang, 2020 (from mid-Cretaceous Burmese amber[21]).

References
  1. Heads, Sam W. (2010). "New Tridactyloidea in Miocene amber from the Dominican Republic (Orthoptera: Caelifera)". Annales de la Société Entomologique de France. N.S. 46 (1–2): 204–210. doi:10.1080/00379271.2010.10697659. ISSN 0037-9271. S2CID 84322530.
  2. Heads, Sam W. (2010). "A new species of Ripipteryx from the Ecuadorian Andes (Orthoptera: Tridactyloidea: Ripipterygidae)". Zootaxa. 2476: 23–29. doi:10.11646/zootaxa.2476.1.3.
  3. Baena-Bejarano, Nathalie J. (2014). "Aspects of the natural history of Ripipteryx (Orthoptera: Ripipterygidae) species in Colombia". Journal of Insect Behavior. 28 (1): 44–54. doi:10.1007/s10905-014-9479-8. ISSN 0892-7553. S2CID 17859291.
  4. Amaral de Gambardella, Loreley (1978). "Bioecología de Rhipiteryx notata Burmeister y su distribución geográfica en el Uruguay (Orthoptera, Tridactyloidea)". Revista de la Sociedad Entomológica Argentina (in Spanish). 37 (1–4). ISSN 1851-7471.
  5. Heads, Sam W.; Taylor, Steven J. (2012). "A new species of Ripipteryx from Belize with a key to the species of the Scrofulosa Group (Orthoptera, Ripipterygidae)". ZooKeys (169): 1–8. doi:10.3897/zookeys.169.2531. ISSN 1313-2970. PMC 3278811. PMID 22371682.
  6. Amaral de Gambardella, Loreley (1971). "Oviposición y eclosión de Rhipipteryx notata Burm. 1838.—(Orthoptera: Tridactyloidea, Rhipipterygidae)". Revista Peruana de Entomología. Anales del Primer Congreso Latinoamericano de Entomología (in Spanish and English). 14 (2): 282–285.
  7. Maes, Jean-Michel (1989). "Catálogo de los Tetrigidae, Rhipipterygidae y Tridactylidae (Orthoptera) de Nicaragua" (PDF). Revista Nicaraguense de Entomología (in Spanish and English). 6: 49–54.
  8. Günther, Kurt K. (1994). "Die Tridactyloidea-Fauna Kolumbiens (Orthoptera, Caelifera)". Deutsche Entomologische Zeitschrift (in German). 41 (1): 1–56. doi:10.1002/mmnd.19940410102.
  9. Song, Hojun; Amédégnato, Christiane; Cigliano, María Marta; Desutter-Grandcolas, Laure; Heads, Sam W.; Huang, Yuan; Otte, Daniel; Whiting, Michael F. (2015). "300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling". Cladistics. 31 (6): 621–651. doi:10.1111/cla.12116. ISSN 0748-3007. S2CID 53702892.
  10. "family Ripipterygidae Ander, 1939". Orthoptera Species File. Retrieved 16 September 2020.
  11. Günther, Kurt (1962). "Eine neue Rhipipteryx-Art aus Ecuador (Orthoptera, Tridactylidae)". Deutsche Entomologische Zeitschrift (in German). 9 (3–4): 251–255. doi:10.1002/mmnd.19620090306.
  12. Günther, Kurt K. (1963). "Über eine Tridactyliden-Ausbeute aus Kolumbien. (Orthoptera)". Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologisches Museum und Institut für Spezielle Zoologie (Berlin) (in German). 39 (1): 187–201. doi:10.1002/mmnz.19630390104.
  13. Günther, Kurt K. (1969). "Revision der Familie Rhipipterygidae Chopard, 1949 (Saltatoria, Insecta)". Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologisches Museum und Institut für Spezielle Zoologie (Berlin) (in German). 45 (2): 259–425. doi:10.1002/mmnz.19690450209.
  14. Günther, Kurt K. (1970). "Rhipipteryx capotensis n. sp., eine neue Art aus Kolumbien. (Rhipipterygidae, Saltatoria, Insecta)". Deutsche Entomologische Zeitschrift (in German). 17 (4–5): 353–356. doi:10.1002/mmnd.19700170406.
  15. Günther, Kurt K. (1976). "Beiträge zur Systematik und Geographischen Verbreitung der Rhipipterygiden (Rhipipterygidae, Tridactyloidea, Saltatoria), Nr. 1". Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologisches Museum und Institut für Spezielle Zoologie (Berlin) (in German). 52 (2): 357–369. doi:10.1002/mmnz.19760520208.
  16. Günther, Kurt K. (1980). "Katalog der Caelifera-Unterordnung Tridactylodea (Insecta)". Deutsche Entomologische Zeitschrift (in German). 27 (1–3): 149–178. doi:10.1002/mmnd.19800270114.
  17. Günther, Kurt K. (1989). "Neue und wenig bekannte Tridactyloidea aus Mittel- und Südamerika (Orthoptera, Caelifera)". Deutsche Entomologische Zeitschrift (in German). 36 (4–5): 347–379. doi:10.1002/mmnd.19890360428.
  18. Baena-Bejarano, Nathalie J.; Heads, Sam W.; Taylor, Steven J. (2018). "Comments on the neglected nymphs of mud crickets in the genus Mirhipipteryx (Caelifera: Tridactyloidea: Ripipterygidae)". Zootaxa. 4486 (2): 180–188. doi:10.11646/zootaxa.4486.2.7. ISSN 1175-5334. PMID 30313760.
  19. Baena-Bejarano, Nathalie J. (2013). Relaciones filogenéticas de los géneros y grupos de especies de Ripipterygidae (Orthoptera: Tridactyloidea) basadas en morfología (M.Sc.) (in Spanish and English). Universidad Nacional de Colombia, Bogotá, Colombia. Retrieved 17 September 2020.
  20. Baena-Bejarano, Nathalie J.; Heads, Sam W. (2015). "Three new species of the genus Ripipteryx from Colombia (Orthoptera, Ripipterygidae)". ZooKeys (502): 129–143. doi:10.3897/zookeys.502.8871. ISSN 1313-2989. PMC 4443593. PMID 26019667.
  21. Xu, Chungpeng; Zhang, Haichun; Jarzembowski, Edmund A.; Fang, Yan (2020). "The first Ripipterygidae (Orthoptera: Caelifera: Tridactyloidea) from mid-Cretaceous Burmese amber". Cretaceous Research. 112: 104356. doi:10.1016/j.cretres.2019.104356.
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