Mantidae

Rivera, Julio, 2010, A historical review of praying mantid taxonomy and systematics in the Neotropical Region: State of knowledge and recent advances (Insecta: Mantodea), Zootaxa 2638, pp. 44-64 : 53-56

Mantidae

Mantidae is the most diverse and variable family, with a wide array of morphological, ecological and behavioural adaptations. As currently conceived, Mantidae is most probably a polyphyletic assemblage ( Svenson & Withing 2004). According to Ehrmann (2002) it includes 20 subfamilies, nine of which occur in the Neotropics.

Angelinae is a quite distinctive group of long, slender-bodied insects, whose species somewhat resemble walking sticks. This subfamily is represented in the Neotropics by two genera. Thespoides Chopard contains a single, very distinctive species from northwestern South America ( T. bolivari Chopard, 1916 ) of which nothing is known besides the original description and a subsequent photo of the holotype by Salazar (2006b). Angela Serville is a widespread genus and well diversified with 17 species. Although Angela is in need of revision, most species are quite distinctive and original descriptions are still useful for the identification of some species. In addition, Lombardo & Agabiti (2001) provided useful descriptions (including male genitalia) of the species occurring in Ecuador: A. armata (De Hann, 1842) , A. guianensis Rehn, 1916 , and A. peruviana (Giglio-Tos, 1916) . This subfamily is sometimes included (possibly incorrectly) within the Thespidae .

Amelinae is a predominantly Old-World assemblage with three genera mainly distributed in the Nearctic region and of which only two occur in the neotropics. The affinities between the New and Old-World two lineages need to be tested. This subfamily consists of small, ground-dwelling species in arid habitats. Litaneutria Saussure includes, according to Beier (1935a), Ehrmann (2002) and Otte & Spearman (2005), seven species. However, Hebard (1935) synonimized most of these species with L. minor (Scudder, 1872) , a highly variable, widely distributed species, and probably the only one that should be included in Litaneutria ; distributional records for this species are found in Hebard (1932) (under L. ocularis Saussure, 1892 ) and Battiston et al. (2005), who also described the male genitalia. Yersinia Saussure, 1869 includes a single species, Y. mexicana ( Saussure, 1859) , occurring in Mexico and Guatemala and whose male genitalia have been depicted by Battiston et al. (2005).

Stagmomantinae consists of three genera mainly distributed in southern North America, Central America and adjacent areas of South America, as well as in the Caribbean region. Phasmomantis Saussure includes two large, rather distinctive species (see Giglio-Tos 1927); Salazar & Stewie (2009) published a photo and information on the holotype of P. championi Saussure & Zehntner, 1894 . Callimantis Stål contains a single insular species widely distributed in the Caribbean region, C. antillarum ( Saussure, 1859) . Lombardo & Perez-Gelabert (2004) depicted this species in detail and discussed its variability and distribution. Stagmomantis Saussure is one of the most diverse genera in the Neotropics, with a range extending well into the Nearctic region. Even though Stagmomantis species are among the most common mantids in Central America, Mexico and the southern U.S., there is not a single comprehensive work dealing with this genus, and the exact number of valid species is uncertain (approximately 20–25 spp.). Although many species are well known and can be identified from the available literature, multiple sources typically must be consulted for accurate identification. Rehn (1935b) is still a good reference for those species occurring in Central America, with detailed descriptions and partial designs of the male genitalia of S. nahua Saussure, 1869 , S. centralis (Giglio-Tos, 1917) , S. vicina ( Saussure, 1870) , S. carolina (Johansson, 1763) , S. montana montana Saussure & Zehntner, 1894 and S. theophila Rehn, 1904 (full genital design of S. theophila in Lombardo & Agabiti 2001). Partial descriptions of the male genitalia of S. hebardi Rehn, 1935 and S. fraterna Saussure & Zehntner, 1894 were made by Rehn (1935a). The male genitalia of S. domingensis (Palisot de Beauvois, 1805) were illustrated by Lombardo & Perez-Gelabert (2004), and those of S. limbata (Hahn, 1835) were depicted by Battiston et al. (2005). Ariza & Salazar (2005) illustrated the male genitalia of Stagmomantis parvidentata colombiana Ariza & Salazar, 2005 . Finally, Salazar (2006b) and Salazar & Stiewe (2009) published photos of the type specimens of (respectively) Stauromantis festae Giglio-Tos, 1917 and Stagmatoptera pagana Saussure, 1870 , both synonyms of S. theophila .

Mellierinae is distributed in southern Mexico, Central America and northern South America, and includes two very similar genera. Xystropeltis Rehn consists of 3 species, although Lombardo (2000b) questioned the placement of X. quadrilobata Terra, 1991 in this genus and suggested that it could be a species of Heterovates Saussure, 1872 (Vatinae) . The other two species ( X. lankesteri Rehn, 1935 and X. meridionalis Lombardo, 2000b ) can be distinguished by consulting the original descriptions ( Rehn 1935a; Lombardo 2000b), but only the male genitalia of X. meridionalis is known ( Lombardo 2000b). Four species are included in Melliera Saussure ; although they are well known in the literature, they need updated revision. Species identifications can be achieved with use of multiple sources ( Rehn 1935a, b, 1951b, 1955).

Stagmatopterinae , a distinctive subfamily within the Neotropical Mantodea , is distributed in tropical America and includes seven genera, none of which have been reviewed in recent years. The last comprehensive revision was by Giglio-Tos (1914), and species identification in this subfamily can be accomplished by consulting this reference in addition to Giglio-Tos (1927). However, given their antiquity, these references often lack enough details and illustrations to confidently separate some of the species. Oxyopsis Caudell consists of 11 poorly known species; the male genitalia have been described for only one species, O. festae Giglio-Tos, 1914 ( Lombardo & Agabiti 2001) . Paroxyopsis Rehn includes a single species, P. i c t e r i c a ( Saussure & Zehntner, 1894), known only from the original description. Catoxyopsis Giglio-Tos also includes a single, uncommon species, C. dubiosa Giglio-Tos, 1898 (female illustrated in Lombardo & Agabiti 2001). Likewise, the five species of Pseudoxyops Saussure & Zehntner also are quite obscure, but a picture of a female of P. perpulchra Westwood was published by Salazar (2003a), and Battiston & Picciau (2008a) described the male and its genitalia, as well as the male genitalia of P. minuta Giglio-Tos, 1914 and P. diluta ( Stoll, 1813) . Parastagmatoptera Saussure includes 13–14 species, but despite this high number it has received exceedingly little taxonomic treatment over the years and is in need of revision. Species recognition by means of the current taxonomic literature is very difficult and several synonymies almost certainly exist. Only the male genitalia of P. unipunctata ( Burmeister, 1838) has been described ( Heitzmann 1959). Lobocneme Rehn is also little known, with just two uncommon species. The male of L. colombiae Hebard, 1919 was pictured by Cerdá (1997a). The well-known genus Stagmatoptera Burmeister includes about 15 species, and despite their commonness and large size, the genus has never been reviewed. Specific identification can be accomplished, within limits, by consulting original descriptions and Giglio-Tos (1914, 1927). The male genitalia of P. biocellata Saussure, 1869 , S. septentrionalis Saussure & Zehntner, 1894 , S. supplicaria ( Stoll, 1813) , S. binotata Scudder, 1896 and S. femoralis Saussure & Zehntner, 1894 were illustrated by Cerdá (1993b), and a SEM image of the male genitalia of S. abdominalis ( Olivier, 1792) was presented by Jantsch (1994b). Ehrmann (1999) published pictures of S. femoralis Saussure & Zehntner, 1894 and S. hyaloptera (Perty, 1832) and established some synonymies. Salazar & Stewie (2009) published a photo and further data on the holotype of S. septentrionalis .

Vatinae is exclusively Neotropical and includes nine genera. Chopardiella Giglio-Tos , formerly placed in Stagmatopterinae and transferred to Vatinae by Ehrmann (2002), has only two species. Lombardo (1994) described the male of C. latipennis ( Chopard, 1912) and included illustrations of the male genitalia. A second species, C. poulaini , is described in detail by Lombardo & Agabiti (2001). Heterovates Saussure , also previously included in Stagmatopterinae , is monospecific; H. pardalina (Saussure, 1872) occurs in northern South America. Its male genitalia were described by Lombardo & Agabiti (2001). Va t e s Burmeister contains13 described species and several undescribed ones. No recent taxonomic revision for this genus is available and the existing literature often is insufficient for reliable species determinations, so old and multiple references must be consulted. Beier (1958) presented a generally useful key to Va t es spp. (but without illustrations) and Lombardo & Agabiti (2001) keyed the species occurring in Ecuador ( V. weyrauchi Beier, 1958 , V. pectinicornis ( Stål, 1877) , V. biplagiata Sjöstedt, 1930 , V. f e s t a e Giglio-Tos, 1898). In this genus only the male genitalia of V. lobata ( Fabricius, 1798) have been described ( Cerdá 1993b). The rare genus Lobovates Deeleman-Reinhold includes a single species, L. chopardi Deeleman-Reinhold, 1957 , known only from two records: the type locality (San Salvador, El Salvador) and Misantla, in Vera Cruz, Mexico ( Lombardo, 2000b, cited as “Misantha), but Lombardo (2000b) also described the male genitalia and Salazar & Stiewe (2009) presented a photograph of the holotype. Hagiotata Saussure & Zehntner consists solely of H. hofmanni ( Saussure & Zehntner, 1894) , a species of southern South America. Phyllovates Kirby includes 14 species of which very little is known. However, Orofino et al. (2006) published a brief study on the species of “western” South America and included descriptions (including male genitalia) of the following species: P. chlorophaea (Blanchard, 1836) , P. brevicornis ( Stål, 1877) , P. s t o l l i ( Saussure & Zehntner, 1894), P. brasiliensis (Piza, 1982b) , P. gracilicollis Orofino et. al, 2006 , P. brevicollis Orofino et. al, 2006 ). In addition, Jantsch (1992a) redescribed the males (including genitalia) of P. inheringi ( Saussure & Zehntner, 1894) and P. brevicornis , and Cerdá (1993b) also illustrated the male genitalia of P. chlorophaea . Pseudovates Saussure is another poorly known taxon of nine species, but it does include the well-known Nearctic species P. arizonae Hebard, 1935 . The only Neotropical species of this genus with recent accounts in the literature is P. peruviana (Rehn, 1911) (see Rivera 2001). The recently described genus Callivates Roy includes a single species from French Guiana, C. stephanei Roy, 2003 . Zoolea Serville includes five species, all treated in an excellent review published by Roy & Ehrmann (2009). A photo of the holotype of Z. lobipes ( Olivier, 1792) is found in Salazar (2006).

Antemninae is a small Central American subfamily with just one described species, Antemna rapax Stål, 1877 ; Salazar & Stiewe (2009) published a photo of the holotype Phyllomantis laurifolia Saussure & Zehntner, 1894 , a synonym of A. rapax . A likely member of this subfamily also occurs in Colombia (Agudelo et al. 2002).

Photinainae is a heterogeneous assemblage of 12 genera, all distributed in the Neotropics. The name Photinainae (formerly Photininae ) was recently amended to avoid homonymy with Photininae (Lampyridae) ( Svenson & Branham 2007). Macromantis Saussure , which includes the largest mantids of the region, consists of four well-defined species ( Roy 2002b). Salazar & Stiewe (2009) published a photo and additional remarks of the holotype of M. nicaraguae Saussure & Zehntner, 1894 . Photina Burmeister includes five poorly known species and is in need of revision. Metriomantis Saussure & Zehntner was partially reviewed by Lombardo (1999), who also illustrated the male genitalia of M. ovata Saussure & Zehntner, 1894 , M. boliviana Lombardo, 1999 and M. occidentalis Lombardo, 1999 . Rivera (2003) described the female of M. boliviana . The female of M. occidentalis was subsequently described by Lombardo & Agabiti (2001), as the female allotype described by Lombardo (1999) was not conspecific with the holotype. The genus Margaromantis Piza includes a single species, M. planicephala ( Rehn, 1916) . The taxonomic and nomenclatural history of this genus and species is complicated, mostly because Margaromantis remained overlooked by taxonomists since its original description (Piza 1982a). Later contributions (Lombardo 1999; Koçak & Kemal 2008; Koçak 2008; Ozdikmen 2008) unwillingly added further confusion. Rivera (2010b) helped to clarify the identity of this genus and its sole species and, as a result, several new synonyms were proposed (see Rivera 2010b for a complete review). Detailed descriptions of M. planicephala are found, as Metriomantis planicephala , in Rehn (1916) and, as Rehniella planicephala , in Lombardo (1999). Photinella Giglio-Tos comprises five poorly known species and is in need of revision; the male genitalia of P. brevis ( Rehn, 1907) has been partially described (Lombardo 1999). Microphotina Beier includes a single species, M. vitripennis (Saussure, 1872) . This genus is also in need of revision since undescribed species certainly occur. Chromatophotina Rivera was recently described and includes two species, C. awajun and C. cofan ( Rivera 2010a) . Hicetia Saussure & Zehntner includes two poorly known species and is in need of revision, as well. Paraphotina Giglio-Tos contains two well-known species and the male genitalia of both species have been described: that of P. reticulata ( Saussure, 1871) by Cerdá (1992) and of P. occidentalis Lombardo, 1998 ( Lombardo 1998) . Orthoderella Giglio-Tos contains two species, O. ornata Giglio-Tos, 1897 and O. delucchii Rivera, 2003 ; male genitalia of both species have been described ( Rivera 2003). Cardioptera Burmeister includes at least five species and was reviewed by Heitzmann-Fontenelle (1968b), with descriptions of the male genitalia. Coptopteryx Saussure is another diverse genus, with about 20 species; it is in need of revision as no recent work has been done on its species. The major references on the taxonomy of Coptopteryx are still Rehn (1913), Giglio-Tos (1915) and Werner (1925). Species identification with these references alone may still be difficult. Zapata (1966) provided a detailed description of C. gayi (Blanchard, 1851) , including the male genitalia, which also were illustrated by Jantsch & Pozza (2001), along with the male genitalia of C. argentina (Burmeister, 1864) . Rivera (2003) illustrated the male genitalia of C. brevipennis Beier, 1858 , and images of both sexes of this species were published in Rivera & Novoa (2006). Brunneria Saussure comprises six species, including B. borealis Scudder, 1869 , a well-known parthenogenetic species with a Nearctic distribution. Very little is known of the Neotropical species and the genus needs revision. Only the male genitalia of B. orinocensis Agudelo & Chica, 2002 is known ( Agudelo & Chica 2002a).

Choeradodinae is comprised of a single genus in the Neotropics, Choeradodis Serville. Roy (2004b) reviewed this genus and provided descriptions of the male genitalia and a key to separate the species. Lombardo (2000a) provided further details on C. rhombicollis (Latreille, 1833) .