______

Proceedings of the Tenth International Symposium on Neuropterology. Piran, Slovenia, 2008. Devetak, D., Lipovšek, S. & Arnett, A.E. (eds). Maribor, Slovenia, 2010. Pp. 269–276.

______

Overview of the taxonomic and biological knowledge of Mexican (Insecta: )

Daniel Reynoso-Velasco1 & Atilano Contreras-Ramos1,2 1Instituto de Biología, Dpto. de Zoología, Apdo. Postal 70-153, 04510 México, D.F., Mexico 2Corresponding autor; E-mail: [email protected]

Abstract. Taxonomic and biological knowledge of Mexican Mantispidae is summarized. Knowledge on this group in Mexico is only fragmentary, particularly in regard to life history. Currently, 23 valid species and three morphospecies have been recorded. Morphospecies of Plega (2) and Trichoscelia (1) probably represent undescribed species. Known species diversity is distributed in the genera Plega (10), Trichoscelia (4), Zeugomantispa (3), (2), Dicromantispa (2), Nolima (2), Entanoneura (1), Leptomantispa (1), and Xeromantispa (1). Few studies treat the of Mexican mantispids. Studies are required for the partially revised Plega and Trichoscelia, while a taxonomic revision of Nolima is close to completion. In regard to biology, it is known that members of Plega and Trichoscelia are predators of some hymenopterans. Mating behavior of one species of Trichoscelia has also been documented.

Key words: Mantispidae, Mexico, taxonomy, biology

Mantidflies are quite particular , because they resemble praying mantises (Mantodea), because of the convergent raptorial forelegs and an elongate, cylindrical prothorax (Reynoso-Velasco & Contreras-Ramos, 2008). The Mantispidae consist of the subfamilies Calomantispinae Navás, Drepanicinae Enderlein, Leach, and Symphrasinae Navás (Lambkin, 1986).

Symphrasinae is distributed from the southwestern United States to Argentina, Drepanicinae occurs in continental Australia and South America, and Calomantispinae in eastern Australia (including Tasmania) and North and Central America, while Mantispinae is considered cosmopolitan occurring between 50° N and 45° S (Lambkin, 1986; Ohl, 2004).

There are few works dealing with Mexican Mantispidae, and those are focused mostly on taxonomy. Only a few publications contain biological notes but there is no comprehensive synthesis of all available data. According to several authors (Penny, 1977; Hoffman, 1992, 2002; Oswald et al., 2002; Ohl, 2004), the Mexican Mantispidae fauna is composed of 23 described species from nine genera and three subfamilies (Table 1). The best known group in Mexico is Mantispinae. Nolima Navás, in Calomantispinae, is undergoing a taxonomic revision, which is close to completion. Other Neotropical genera in Symphrasinae, such as Plega Navás and Trichoscelia Westwood, require taxonomic revision, and when this is achieved, taxonomic knowledge on the Mexican mantispid fauna will be in an acceptable state.

Overall, the biology of Mantispidae is poorly understood. Larvae of Mantispinae are obligate predators on spiders egg sacs; Calomantispinae larvae feed on a wide variety of sedentary prey under laboratory conditions; and larvae of Symphrasinae have been found in association with insect immature stages. There is no information on the biology of Drepanicinae (Redborg, 1998). In this paper, we present a summary of available information about the taxonomy and biology of Mexican Mantispidae based on literature, in order to foster knowledge on this insect group. The term biology is used broadly, to mean studies related to natural history, life history, and

ISBN 978-961-6657-16-7 © 2010 Faculty of Natural Sciences and Mathematics UM other non taxonomic subjects. Would there be formal ecological studies, or of other sort (e.g., physiology, genetics), these had been included, but so far no such studies have been performed on Mexican fauna. As part of the taxonomic knowledge, distribution by state of every species is listed and illustrated.

Results

Taxonomic knowledge

According to previous knowledge, the Mexican Mantispidae fauna has representatives from three subfamilies, nine genera, and 23 species. Reynoso-Velasco & Contreras-Ramos (2008) estimated a fauna of 28 species, but this evaluation contained five morphospecies, of which only three might actually represent undescribed species (Table 1).

Subfamily Calomantispinae Navás This subfamily is represented in Mexico by the genus Nolima Navás. Since the description of Nolima, N. victor Navás and N. praeliator Navás were considered to be present in Mexico (Navás, 1914b). Reynoso-Velasco & Contreras-Ramos (2008) cited the occurrence of both species and also two morphospecies. At this moment, we know that only two species of Nolima occur in Mexico, within the Neotropical area of the country (Table 1), with records from six states (Figs. 1 and 2).

Subfamily Mantispinae Leach This subfamily is represented in Mexico by ten species in six genera (Table 1). This information is quite precise and reliable due to Hoffman’s work (1992, 2002) on the systematics of the Mantispinae of the New World. These two works by Hoffman are excellent sources to identify the species of this subfamily (Table 2). There are records of Mantispinae from 27 states over the entire country (Figs. 17–26). In this group, five species can be considered widespread in Mexico (Figs. 17, 19, 20, 22, and 25).

Subfamily Symphrasinae Navás This subfamily is represented in Mexico by 11 nominal species and three morphospecies (Table 1), which probably represent undescribed species. Both Plega Navás and Trichoscelia Westwood lack taxonomic revisions, and because of that, the exact number of species is not known. There could be synonyms as well as undescribed species in both genera. Currently, it is difficult to identify specimens in these genera, as only the original species descriptions are available (Table 2). The subfamily is present in both the Nearctic and Neotropical areas of the country. There are records from 17 states (Figs. 3–16), although there are some species only known from the type locality (Table 1).

Biological knowledge

Overall, Mantispinae is the best known group in the New World, both in systematics and biology. Nonetheless, regarding mantispid natural history of Mexican species, this is not the case, because there is only information on the Symphrasinae.

In the case of Plega melitomae Linsley and MacSwain, it was found that immatures of this species are predators of immature stages of the bee Melitoma euglossoides Lepeletier and Serville, because some mantispids from this species were reared from the bee’s nest cells (Linsley & MacSwain, 1955).

Plega fumosa Linsley and MacSwain has been considered to be associated (potential predators) with paper wasps, as adults of these mantispids were collected from an Acacia-like legume plant, where three species of Polistes were nesting (Linsley & MacSwain, 1955).

Larvae from Plega yucatanae Parker and Stange have been collected in association with immatures of Megachile exaltata Smith. Newly emerged adult mantispids were collected from the top of a cave where there were bee nests; after checking the nest cells, the mantispid immatures were found (Parker & Stange, 1965).

About Trichoscelia santareni (Navás), there is information concerning the reproductive behavior and posterior invasion of a wasp colony pertaining to the species Polybia diguetana Buysson (Dejean & Canard, 1990). The process began with males and females flying and forming a cloud, after that the male started an approach flight

270 Table 1. Mantispidae species present in Mexico by state (modified from Reynoso-Velasco, 2007). Species marked with an asterisk (*) are known only from the type locality record.

Taxon State

Symphrasinae Plega banksi Rehn Chihuahua, Sinaloa, Sonora Plega dactylota Rehn Baja California Sur, Chihuahua, Coahuila, Jalisco, Sinaloa, Sonora Plega fratercula Rehn Baja California Sur, Chihuahua *Plega fumosa Linsley and MacSwain Michoacán Plega melitomae Linsley and MacSwain Chiapas Plega signata (Hagen) Baja California, Baja California Sur Plega variegata Navás Puebla, Veracruz Plega yucatanae Parker and Stange Chiapas, Yucatán Plega sp. 1 Guerrero, Jalisco, Morelos, Nayarit, Veracruz *Plega sp. 2 Sonora *Trichoscelia banksi Enderlein Chiapas Trichoscelia santareni (Navás) Quintana Roo, Tabasco *Trichoscelia tobari (Navás) Tabasco *Trichoscelia sp. 1 Sonora

Calomantispinae Nolima infensus Navás Morelos, Oaxaca Nolima victor Navás Chiapas, Guerrero, Hidalgo, Oaxaca, Querétaro

Mantispinae (Say) Baja California Sur, Chiapas, Chihuahua, Colima, Durango, Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Sinaloa, Tabasco, Tamaulipas, Veracruz Climaciella semihyalina (Le Peletier de Veracruz Saint-Fargeau and Audinet-Serville) Dicromantispa interrupta (Say) Baja California, Baja California Sur, Chiapas, Chihuahua, Coahuila, Colima (Revillagigedo islands), Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, San Luis Potosí, Sinaloa, Sonora, Tamaulipas, Veracruz, Yucatán Dicromantispa sayi (Banks) Campeche, Chiapas, Chihuahua, Durango, Guerrero, Nuevo León, Quintana Roo, Sinaloa, Tabasco, Tamaulipas, Veracruz Entanoneura batesella (Westwood) Chiapas, Jalisco, Oaxaca Leptomantispa pulchella (Banks) Baja California Sur, Chiapas, Chihuahua, Hidalgo, Jalisco, Michoacán, Nuevo León, Oaxaca, San Luis Potosí, Sinaloa, Tamaulipas, Veracruz Xeromantispa scabrosa (Banks) Nuevo León Zeugomantispa compellens (Walker) Campeche, Chiapas, Oaxaca, Quintana Roo, San Luis Potosí, Veracruz Zeugomantispa minuta (Fabricius) Campeche, Chiapas, Chihuahua, Colima, Guerrero, Hidalgo, Jalisco, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Sinaloa, Tabasco, Tamaulipas, Veracruz, Yucatán Zeugomantispa virescens (Rambur) Campeche, Chiapas, Colima, Jalisco, Nayarit, Oaxaca, Quintana Roo, San Luis Potosí, Veracruz

towards the motionless female and initiated a courting behavior by moving the antennae and the forelegs; posteriorly, they clasped their genitalia and maintained this position for about 40–60 minutes. The entire sequence took 149 minutes. At the end, the male of the mantispid flew away while the female groomed her genitalia, and finally she flew towards the wasp nest. Dejean & Canard (1990) also suggested that it is possible that the female from this species produces a tolerance allomone that permits females to enter into the nest avoiding the guards at the entrance.

271 Figs. 1–2. Known species distribution of Calomantispinae Navás in Mexico depicted by state.

Table 2. Taxonomic references for Mexican species of Mantispidae (Reynoso-Velasco & Contreras-Ramos, 2008).

Taxon Selected references

Calomantispinae Navás Navás (1914b), Rehn (1939) Mantispinae Leach Hoffman (1992, 2002) Symphrasinae Navás Enderlein (1910), Navás (1914a, 1928), Rehn (1939), Linsley & MacSwain (1955), Parker & Stange (1965), Penny (1982), Rice (1987)

Discussion

In general, the taxonomic knowledge about Mexican Mantispidae may be considered fairly complete. Considering only numbers of valid species, it could be stated that most species of Mantispinae are known, and the same happens in Calomantispinae now that the taxonomic revision of Nolima Navás is nearly finished. However, there is taxonomic effort needed in the case of Symphrasinae, as both Plega Navás and Trichoscelia Westwood still need a comprehensive taxonomic revision. Currently, the best estimation on the Mexican Mantispidae species richness is 26 species (including three morphospecies), two from Calomantispinae Navás, 14 from Symphrasinae Navás, and 10 from Mantispinae Leach. In summary, most Mexican mantispid species may be identified with no problem, with the exception of Symphrasinae (Table 2).

Regarding distributional knowledge in Mexico, this is based on scattered records. There has never been a systematic collecting effort on this insect group, and because of this the information is fragmentary and taxonomically uneven. There are some species of Symphrasinae just known from the type locality, and five states (Aguascalientes, Distrito Federal, Guanajuato, Tlaxcala, and Zacatecas) lack mantispid distributional records at all. The best known geographical distribution is from Mantispinae, which may reflect they are more common when collecting. At this time, it is difficult to talk about distribution patterns in Mexico, although it is clear that there are some widespread species in Mantispinae and some possibly endemic species belonging to Symphrasinae.

In the biological aspect, the information comes from four incidental encounters with Symphrasinae species in Mexico, but even so, these studies constitute important information sources about feeding habits and reproductive behavior of these insects, not only for the Mexican fauna. Of course, there is a need for work especially focused on mantispids in the field and laboratory conditions in order to learn more about mantispid biology.

Acknowledgements. One of us (DRV) wants to thank to the Posgrado en Ciencias Biológicas, UNAM for support for his studies.

272

Figs. 3–10. Known species distribution of Symphrasinae Navás in Mexico depicted by state.

273

Figs. 11–16. Known species distribution of Symphrasinae Navás in Mexico depicted by state. Figs. 17–18. Known species distribution of Mantispinae Leach in Mexico depicted by state.

274

Figs. 19–26. Known species distribution of Mantispinae Leach in Mexico depicted by state.

275

References

Dejean, A. & Canard, M. 1990. Reproductive behavior of Trichoscelia santareni (Navás) (Neuroptera: Mantispidae) and parasitization of the colonies of Polibya diguetana R. du Buysson (Hymenoptera: Vespidae). Neuroptera International, 6: 19–26.

Enderlein, G. 1910. Klassifikation der mantispiden nach dem Material des Stettiner Zoologischen Museums. Stett. Entomol. Zeit., 71: 341–379.

Hoffman, K.M. 1992. Systematics of the Mantispinae (Neuroptera: Mantispidae) of North, Central, and South America. Ph. D. dissertation, Clemson University, Clemson, S.C.

Hoffman, K.M. 2002. Family Mantispidae, pp. 251–275. In: Penny, N. D. (ed.), A guide to the lacewings (Neuroptera) of Costa Rica. Proc. Cal. Acad. Sci., 53: 161–457.

Lambkin, K.J. 1986. A revision of the Australian Mantispidae (Insecta: Neuroptera) with a contribution to the classification of the family. I. General and Drepanicinae. Aust. J. Zool., Suppl. Ser., 116: 1–142.

Linsley, E.G. & MacSwain, J.W. 1955. Two new species of Plega from Mexico (Neuroptera: Mantispidae). Pan-Pac. Entomol., 31: 15–19.

Navás, L. 1914a. Névroptères nouveaux de l’Amerique du Nord. Iième série. Entomol. Zeit., Frankfurt am Main, 28: 18–20, 25–26.

Navás, L. 1914b. Mantíspidos nuevos. 2. a serie. Mem. R. Acad. Cienc. Artes Barc., 11: 83–103.

Navás, L. 1928. Insectos neotrópicos. 3.a serie. Rev. Chil. Hist. Nat., 31: 316–328.

Ohl, M. 2004. Annotated catalog of the Mantispidae of the World (Neuroptera). Contr. Entomol. Int., 5: 131–262.

Oswald, J., Contreras-Ramos, A. & Penny, N.D. 2002. Neuroptera (Neuropterida). In: Llorente, B.J. & Morrone, J.J. (eds.), Biodiversidad, taxonomía y biogeografía de artrópodos de México: Hacia una síntesis de su conocimiento. Vol. III. Universidad Nacional Autónoma de México, Mexico City. Pp. 559–581.

Parker, F.D. & Stange, L.A. 1965. Systematic and biological notes on the tribe Platymantispini (Neuroptera, Mantispidae) and the description of a new species of Plega from Mexico. Can. Entomol., 97: 604–612.

Penny, N. D. 1977. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, ilhas Caraíbas e América do Sul. Acta Amazonica, 7: 1–61.

Penny, N. D. 1982. Neuroptera of the Amazon basin. Part 6. Mantispidae. Acta Amazonica, 12: 415–463.

Redborg, K. E. 1998. Biology of the Mantispidae. Annu. Rev. Entomol., 43: 175–194.

Rehn, J.W.H. 1939. Studies in North American Mantispidae (Neuroptera). Trans. Am. Entomol. Soc., 65: 237–264.

Reynoso-Velasco, D. 2007. Mantispidae (Insecta: Neuroptera) de México. In: Estrada, E., Equihua, A., Luna, C. & Rosas, J. L. (eds.), Entomología Mexicana. Vol. 6. Sociedad Mexicana de Entomología A.C. Mexico. Pp. 1408–1412.

Reynoso-Velasco, D. & Contreras-Ramos, A. 2008. Mantispidae (Neuroptera) of Mexico: distribution and key to genera. Ann. Entomol. Soc. Am., 101: 703–712.

Rice, M. E. 1987. Morphological variation in Plega dactylota (Neuroptera: Mantispidae) and status of its subspecies. J. Kans. Entomol. Soc., 60: 341–344.

276