Liturgusa maya Saussure & Zehntner, 1894

Svenson, Gavin J., 2014, Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini), ZooKeys 390, pp. 1-214 : 59-64

Liturgusa maya Saussure & Zehntner, 1894

Mantis annulipes (non Mantis annulipes Audinet Serville, 1838): von Charpentier 1843: 28-29, pl. 28 (partim).

Liturgousa annulipes : Saussure 1871: 102 (♂ only, partim).

Liturgousa cayennensis [Var.] maya: Saussure and Zehntner 1894: 160; Scudder 1901: 159, 407; Rehn 1903: 6; Marshall 1975: 318; Roy and Cuche 2008: 14, 21.

Liturgousa maya : Scudder 1901: 159, 419; Kirby 1904: 271; Hebard 1932: 211; Rehn 1935: 201, pl. 8, fig. 5; Hughes-Schrader 1950: 11-14, 38, 45, Table 1, Fig. 9; Hughes-Schrader 1951: 178, 180, 183-184, 186-187, Tables 1-2, Fig. 1; Hughes-Schrader 1953: 544-554; Henderson 1965: 215; Cerdá 1996: 76;

Liturgusa maya : Giglio-Tos 1927: 293; Beier 1935: 11; Jantsch 1991: 125; Terra 1995: 54; Jantsch 1999: 48; Maes and Roy 2000: 61; Lombardo and Agabiti 2001: 90, 96; Ehrmann 2002: 207; Agudelo 2004: 55, Table 3.1; Agudelo 2005: 3; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116, 142; Svenson and Whiting 2009: Appendix S1.

Liturgusa maja : Passerin d‘Entrèves 1981: 61.

Liturgusa charpentieri : Giglio-Tos 1927: 294; Beier 1935: 11; Passerin d‘Entrèves 1981: 61; Terra 1995: 53; Salazar E. 1998: 105, Fig. 4; Jantsch 1999: 48; Salazar E. 1999: 10; Lombardo and Agabiti 2001: 90; Salazar E. 2002: 124; Ehrmann 2002: 207; Agudelo 2004: 55, Table 3.1; Agudelo 2005: 3; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116, 141. syn. n.

Type.

Lectotype Male. The Natural History Museum (British Museum Natural History), London, UK.

Type locality.

Temax, N. Yucatan, Gaumer (Lat. 21.143702, Long. -88.942315).

Material examined.

Liturgusa maya Saussure & Zehntner, 1894.

Taxonomic history.

One of the earliest species to be described, Liturgusa maya was referenced in early works as Mantis annulipes , but this is likely due to the lack of comparison with the type of Mantis annulipes . Saussure and Zehntner recognized the species as unique, but as a variant of Liturgusa cayennensis . The species was later formalized as being distinct. It is apparent from the references that include Liturgusa maya , the species name was largely used as a default species identification. This is likely due to the limited state of knowledge surrounding the species boundaries within the genus. Therefore, many of the previous records of Liturgusa maya cannot be accurately confirmed or falsified and using historical records for biogeographic studies or population occurrence studies should be avoided.

Five syntypes designated by Saussure and Zehntner were examined from The Natural History Museum, London, and Muséum d’Histoire naturelle, Geneva. To increase taxonomic stability within the species, under Article 74.1 of the International Code of Zoological Nomenclature a male syntype from the BMNH has been selected to become the unique bearer of the name of the nominal species-group Liturgusa maya (lectotype). Two syntype females from the BMNH and one additional syntype male and one additional syntype female from the MHNG all become paralectotypes under Article 74.1.3 of the Code.

An extensive search for the holotype of Liturgusa charpentieri was conducted in collections in Italy, but was never located (see Taxonomic History of Liturgusa guyanensis ). However, the original description given by Giglio-Tos (1927) for Liturgusa charpentieri matches Liturgusa maya and with the expanded range of Liturgusa maya into South America as evidenced by specimens linking Central and South America through occurrences in Colombia, Venezuela, Ecuador and Peru, Liturgusa charpentieri can no longer be considered as distinct based on its southern distribution. Therefore, Liturgusa charpentieri Giglio-Tos, 1927, is now considered as a junior synonym to Liturgusa maya Saussure & Zehntner, 1894.

Interestingly, Liturgusa maya was included in a number of studies focused on chromosomes headed by Sally Hughes-Schrader in the 1940's and into the 60's.

Natural history.

The species has been found in wet tropical forests in Nicaragua, Costa Rica, Peru, and numerous other regions within its range. However, individuals have also been found in seasonally dry forests, open habitats as well as heavily impacted habitats such as park-land or the edges of parking lots. Perhaps the versatility of Liturgusa maya in habitat utilization has led to its broad geographical range that extends from mid-Mexico to southern Peru. The current thinking for why Liturgusa is so diverse relates to the narrow geographic ranges of species that is linked to poor dispersal ability. However, Liturgusa maya appears to violate this thinking. Although genitalic and external morphology are rather consistent across the entire range, genetic information may lead to the discovery of considerable new diversity by uncovering cryptic species. Without an accurate model of Liturgusa species origins, it can not be said whether Liturgusa maya is young or old relative to other Liturgusa species. Perhaps the species is undergoing a dispersal and speciation event that genetic information could uncover. It is suggested that a population genetics study of Liturgusa maya would be the logical next step to understanding how the species could have such an amazing geographic and habitat range.

As with most Liturgusa species, Liturgusa maya are adept runners and live on medium size smooth-bark tree trunks. Since they are found in wet and dry forests, some individuals have been observed on a broader diversity of tree types, some with moss or even rough bark. Size variation within Liturgusa maya is more extreme than any other species of Liturgusa . The largest female is 145% the size of the smallest, a disparity not matched in the rest of Liturgusa (139.5% for Liturgusa nubeculosa , which is a much larger species).

Diagnosis.

Most similar to Liturgusa kirtlandi and Liturgusa trinidadensis with a similar size, coloration and pronotum shape, Liturgusa maya is distinct from the other two by a number of features including male genitalia. The most obvious difference easily diagnosing Liturgusa maya from Liturgusa kirtlandi is that the apical process (paa) is elongate, thickened and the terminus forms an evenly rounded terminus rather than an angled blunt tip. In addition, Liturgusa maya can be differentiated from Liturgusa trinidadensis by the larger apofisis falloid (afa) compared to the barely present, but sharp structure seen in Liturgusa trinidadensis . The species has the greatest size variation across its range. The main difference between Liturgusa maya and Liturgusa kirtlandi is the presence of tubercles on the pronotum in Liturgusa kirtlandi .

Description.

Male. (Figs 13A, 14A) N=18: Body length 19.38-25.46 (22.43); forewing length 13.16-16.56 (14.87); hindwing length 10.69-13.58 (12.28); pronotum length 5.61-7.39 (6.30); prozone length 1.68-2.36 (1.95); pronotum width 2.11-2.99 (2.44); pronotum narrow width 1.57-2.14 (1.77); head width 4.46-6.34 (4.99); head vertex to clypeus 1.79-2.67 (1.98); frons width 1.61-2.39 (1.80); frons height 0.61-0.93 (0.71); prothoracic femur length 5.40-7.42 (6.25); mesothoracic femur length 3.32-8.28 (7.34); mesothoracic tibia length 5.22-7.36 (6.10); mesothoracic tarsus length 4.58-8.07 (5.55); metathoracic femur length 6.61-9.26 (7.86); metathoracic tibia length 7.35-10.45 (8.69); metathoracic tarsus length 6.70-8.69 (7.74); pronotal elongation measure 0.30-0.33 (0.31); pronotal shape measure 0.37-0.41 (0.39); head shape measure 0.37-0.42 (0.40); frons shape measure 0.36-0.42 (0.39); anteroventral femoral spine count 14-16 (15); anteroventral tibial spine count 9-11 (10); posteroventral tibial spine count 7.

Head (Fig. 42A): Transverse, the juxta-ocular protuberances small, but pronounced, the apex just lateral to the midline; the vertex is straight, but sometimes dips just prior to the parietal sutures, even with the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc, the entire carina depressed into the head. Ocelli small, the central more enlarged (about twice the size as the lateral), all protruding on small cuticular mounds; the lateral ocelli oriented outward. The carina on the frons not very pronounced, the medial region just ventral to the carina depressed. Clypeus transverse, the upper margin slightly convex, the lower margin slightly concave or straight; the central, transverse carina pronounced and straight. Antennae scape and pedicel pale, the flagellum black just slightly distal to the base. Black band extending straight over the medial carina of the frontal suture, the very center of the carina pale; black markings extend ventrally and dorsally from black band; two prominent pale marks positioned just lateral to the lateral ocelli; two pale marks positioned on the lower region of the vertex. Lower region of frons darkly pigmented; the dorsal half of clypeus darkly pigmented, the ventral half pale; the mandibles and labrum with pale and brown markings; the vertex and juxta-ocular protuberances mostly black with pale speckles; the area immediately adjacent to lateral ocelli black. Palpi are pale.

Pronotum (Fig. 47Q): About three times long as wide with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone square with slightly convex margins that gradually taper to an evenly rounded anterior margin; margins smooth or with very few blunt tubercles. Metazone with concave lateral margins without interruptions or bulges; margins with small tubercles; posterior margin with a medial emargination; the dorsal surface of the posterior third of the metazone slightly depressed. Mostly dark with pale and black marking across the surface, faint swirls present on the metazone just posterior to the supra-coxal sulcus.

Prothoracic Legs: Femur robust with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a black band running medially from the base to terminus that may be interrupted, various black marks present in addition to the band that correspond to banding marks on the posterior surface; the ventral surface pale. Posterior surface of femur with few tubercles. A femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and exactly between the first two proximal posteroventral spines, in line with the most distal discoidal spine; pit is pigmented darkly. Posterior prothoracic femoral genicular spine smaller than posteroventral spines (highly variable), originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the third through sixth of similar length, the second slightly longer. Prothoracic coxae smooth, the anterior surface with a very small, black mark medially in the proximal half as well as a very small black spot medially towards the distal terminus.

Meso- and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina present. Mesotarsi with first segment as long or slightly longer than the remaining segments combined.

Wings: Forewings mottled with brown, pale and greenish coloration; the costal region with defined banding, green and brown alternating markings, the brown marks smaller; vein coloration mostly corresponding with surrounding colors; two pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein; a large pale area is positioned centrally. Forewings often, but not always asymmetrically colored; one being mottled as described the other is darkened significantly with a black or rust tone, the mottled pattern still visible; extending just beyond or as long as the abdomen. Hindwings with opaque discoidal region, colored rust proximally and along the anterior margin, otherwise black; the anal region smoky black and translucent; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing elongate.

Abdomen: Slightly widened in the middle, the fourth tergite the widest region before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.

Genital Complex (Fig. 51I.1-I.4): The main body of ventral left sclerite (L4A) with rounded terminus, but with a distal process (pda) positioned just lateral to the midline that is rounded (could be short and rather blunt or more elongate and narrow), projecting at an angle, appearing like a small, well-sclerotized tooth; sometimes a depression on the opposite lateral half from the pda is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) short, quickly narrowing to a sharp point, sometimes curved and sometimes with a rough surface; the apical process (paa) elongate and thin, the terminus an evenly rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus; the ventral plate (pia) long, broad proximally with strongly defined grooves; the ventral process (pva) tooth-like and curved at the proximal base, the distal tip narrowing with a rapid constriction towards the end.

Redescription.

Female. (Figs 12B, 13B, 14C) N=28: Body length 24.02-33.46 (27.24); forewing length 14.55-21.12 (16.95); hindwing length 12.02-15.61 (13.82); pronotum length 6.75-8.99 (7.46); prozone length 2.08-2.83 (2.31); pronotum width 2.74-3.55 (2.97); pronotum narrow width 1.89-2.57 (2.16); head width 5.26-6.94 (5.94); head vertex to clypeus 2.20-2.88 (2.49); frons width 2.03-2.75 (2.30); frons height 0.76-0.94 (0.85); prothoracic femur length 6.73-8.48 (7.31); mesothoracic femur length 7.64-10.25 (8.55); mesothoracic tibia length 5.74-8.19 (6.77); mesothoracic tarsus length 4.57-7.66 (5.9); metathoracic femur length 7.76-10.13 (8.53); metathoracic tibia length 8.41-11.21 (9.68); metathoracic tarsus length 7.38-11.03 (8.53); pronotal elongation measure 0.30-0.33 (0.31); pronotal shape measure 0.38-0.41 (0.40); head shape measure 0.38-0.46 (0.42); frons shape measure 0.34 -0.41 (0.37); anteroventral femoral spine count 14-16 (15); anteroventral tibial spine count 7-10 (10); posteroventral tibial spine count 7.

Head (Fig. 42B): About as broad as long, the juxta-ocular protuberances large; the vertex higher than the dorsal margin of the eyes.

Pronotum (Fig. 47R): The dorsal surface of the posterior third of the metazone not depressed.

Prothoracic Legs: As described for males.

Meso- and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined.

Wings: The costal region of forewing with less defined banding, proximal region mostly mottled with brown and pale. Forewings extending just shy of the terminus of the abdomen, the last few segments and supra-anal plate mostly visible.

Abdomen: Widened, the fifth tergite the widest region before a gradual posterior narrowing. Tergites without posterolateral tergal projections. Supra-anal plate about as long as wide, rounded terminus.