Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini)

Abstract The praying mantis genus Liturgusa Saussure, 1869 occurs only in Central and South America and represents the most diverse genus of Neotropical Liturgusini (Ehrmann 2002). The genus includes bark dwelling species, which live entirely on the trunks and branches of trees and run extremely fast. All species included within the genus Liturgusa are comprehensively revised with a distribution stretching from central Mexico, the island of Dominica to the southeastern regions of Brazil and southern Bolivia. All known species are redescribed to meet the standards of the new treatment of the genus (11 species). Three new genera are described including Fuga gen. n., Velox gen. n., and Corticomantis gen. n. for species previously included in Liturgusa as well as Hagiomantis. Liturgusa mesopoda Westwood, 1889 is moved to within the previously described genus Hagiomantis Audinet Serville, 1838. A total of 19 species are newly described within Liturgusa, Fuga, and Velox including L. algorei sp. n., L. bororum sp. n., L. cameroni sp. n., L. cura sp. n., L. dominica sp. n., L. fossetti sp. n., L. kirtlandi sp. n., L. krattorum sp. n., L. manausensis sp. n., L. maroni sp. n., L. milleri sp. n., L. neblina sp. n., L. purus sp. n., L. stiewei sp. n., L. tessae sp. n., L. trinidadensis sp. n., L. zoae sp. n., F. grimaldii sp. n., and V. wielandi sp. n. Four species names are synonymized: Liturgusa peruviana Giglio-Tos, 1914, syn. n. = Liturgusa nubeculosa Gerstaecker, 1889 and Hagiomantis parva Piza, 1966, syn. n., Liturgusa sinvalnetoi Piza, 1982, syn. n., and Liturgusa parva Giglio-Tos, 1914, syn. n. = Mantis annulipes Audinet Serville, 1838. Lectotypes are designated for the following two species: Liturgusa maya Saussure & Zehntner, 1894 and Fuga annulipes (Audinet Serville, 1838). A male neotype is designated for Liturgusa guyanensis La Greca, 1939. Males for eight species are described for the first time including Liturgusa cayennensis Saussure, 1869, Liturgusa lichenalis Gerstaecker, 1889, Liturgusa guyanensis La Greca, 1939, Liturgusa maya Saussure & Zehntner, 1894, Liturgusa nubeculosa Gerstaecker, 1889, Fuga annulipes (Audinet Serville, 1838), Corticomantis atricoxata (Beier, 1931), and Hagiomantis mesopoda (Westwood, 1889). The female of Fuga fluminensis (Piza, 1965) is described for the first time. Complete bibliographic histories are provided for previously described species. The spelling confusion surrounding Liturgusa/Liturgousa is resolved. Full habitus images for males and females are provided for nearly all species. Habitus and label images of type specimens are provided when possible. Diagnostic illustrations of the head and pronotum for males and females are provided for all species when possible. Illustrations of male genital structures are provided for all species for which males are known. Measurement data, including ranges and averages, are provided for males and females of all species. Combined male and female genus and species level dichotomous keys are provided with a Spanish translation. A complete table of all examined specimens lists label data, museum codes, repositories, and other specimen specific information. A KML file with all georeferenced locality records is downloadable from mantodearesearch.com for viewing in Google Earth. Natural history information is provided for species observed by the author.


Introduction
The family Liturgusidae (sensu Ehrmann 2002) includes a broad assemblage of genera distributed on five continents, all members being characterized as ecomorphic specialists on tree trunks or branches. Informally called "bark mantises", the group exhibits: heavy camouflage mottling that includes browns, black, and other earth tone colors matching tree bark or lichen substrates; dorsoventral flattening for a lower profile against a flat surface; and a ventral prothoracic femoral pit to accommodate for the distal posteroventral prothoracic tibial spine (homoplasious with other Mantodea, see Wieland 2013). However, the family, which appears to be based on a strong ecomorphic grouping, was found to be polyphyletic by Svenson and Whiting (2009) who used molecular data to reconstruct the phylogeny. They found that bark mantis genera group more closely to other, morphologically dissimilar taxa distributed within the same geographic region. This led to the conclusion that higher-level classification of Mantodea was largely confounded by morphological convergence based on similar ecomorphic adaptations in independent lineages. In addition, Wieland (2013) found the family to be non-monophyletic within a thorough morphological phylogenetic analysis.
The Neotropical Liturgusidae (sensu Ehrmann 2002) includes three described genera and 24 species that are distributed from the southern United States and a few Caribbean Islands to the southern regions of Brazil and Bolivia. Liturgusa Saussure, 1869, Hagiomantis Audinet Serville, 1838, and Gonatista Saussure, 1869 are all distinct genera, the first two forming a monophyletic lineage while Gonatista was not included in the phylogeny (Svenson and Whiting 2009). However, recently generated molecular data from a species of Gonatista from the Dominican Republic places the genus outside of Neotropical Liturgusidae (Rivera and Svenson, unpublished data). This study focuses primarily on the species included within Liturgusa, but the type specimens of the highly similar Hagiomantis species were examined to ensure species level placements within these two genera were correct.

Taxonomic history
Jean-Guillaume Audinet Serville (1838: 199) was first to describe a Neotropical Liturgusini species with the description of Mantis annulipes, which De Haan later included within the Oxypilus subgroup within Mantis (De Haan 1842: 84).
More than 30 years later, Henri Louis Frédéric de Saussure (1869: 62) described another species, L. cayennensis, and created a new genus, Liturgousa, for his new species as well as M. annulipes. Saussure (1872: 260) later described L. surinamensis . Carl Stål (1877: 3, 40) changed the spelling of Saussure's genus to Liturgusa, which is now the spelling of prevailing use (see Taxonomic History under the genus description below for nomenclature information).

Natural history
Members of the Neotropical Liturgusini are strictly associated with tree bark habitats, showing preference for smooth bark, presumably for ease of running. However, specimens have been found on trees with mossy covering across as much as fifty percent of the trunk.
Individuals are typically found with the long axis of the body aligned with the vertical axis of the tree in a head down position ( Fig. 1A-C). The posterior portions of the abdomen touch or nearly touch the bark while the head and prothoracic legs are held in an elevated position, the body angled relative to the trunk. Flight is rarely observed in all species of Neotropical Liturgusini, but males of L. fossetti sp. n. found in Costa Rica have been observed flying to nocturnal light traps (Lord personal observation), indicating some species may have more active flight in males than in females.
Being highly visual predators like other Mantodea, individuals easily detect approaching people and will run laterally to the opposite side of the tree when approached during daylight hours, although many have been observed assuming a crouching position before bolting. Individuals approached at night are far less prone to run, possibly due to low light conditions or more likely the effect of bright lights from flashlights obscuring or saturating their vision. If pursued, individuals run up the tree at a continuous gate or in spurts, but ultimately achieve unreachable heights. Although their exact speeds are not known, they run incredibly fast. Their speed and cursorial lifestyles on flat, but vertical tree trunks is reminiscent of ground hunting tiger beetles, a species of which, Cicindela repanda Dejean, 1825, was well characterized by Gilbert (1997). It would be interesting to conduct similar work on a species of Liturgusa.
Although diurnal capture is possible, nocturnal collecting proved to be far more productive. Using a spotlight or head lamp along with a large vial (50 ml), one can approach from a distance while shining the light on the mantis. They typically don't move in this circumstance, which allows enough time to position the vial at an angle anterior to the head and either swiftly brought down over the specimen or with a touch to their posterior they run forward and inside the vial. Diurnal capture can be successful by searching trees with an insect net or curved branch, tapping the reverse side of trunks since they often have fled to the other side before being spotted. Tapping the reverse side of the trunk will drive them around to the visible side of the tree. By corralling individuals with hands and arms, which typically results in hugging trees, to lower positions on the trunk, a quick movement of a cupped hand over the specimen will achieve capture. On occasion, individuals will escape to the ground cover at the base of the tree, running rapidly and being still to avoid detection. Rarely, species have been observed taking flight directly off the tree and flying a moderate distance before dropping onto a plant or hitting the forest floor (males of L. algorei sp. n. in Peru). Larger females of L. maya have been observed to take flight, flutter a short distance and drop into leaf litter and exhibit thanatosis.
Most individuals are collected in the lower sections of tree trunks, but this may be mostly related to the limited searching ability of the collector that is walking through  Rehn, 1935, female from Nicaragua B L. kirtlandi sp. n., female from Bolivia C L. nubeculosa Gerstaecker, 1889, female from Bolivia D ootheca of unknown species of Liturgusa. the forest. Specimens have been observed in higher regions of tree (L. nubeculosa, L. fossetti sp. n., and L. krattorum sp. n.) as well as within the crown of trees (L. nubeculosa) through canopy rope accession. Species are also plentiful in canopy fogging efforts like those performed by Terry Erwin in southern Peru where a large number of Liturgusa were sampled (L. lichenalis). It is not known if bark mantises have a preference for their vertical position on a tree or how extensively they use more terminal branches higher in the canopy since this is obviously difficult to observe, but would be interesting to research.
Although empirical data is lacking, a general trend of small species preferring smaller trunk diameters while larger species are typically more common on large trunked trees has been observed in Bolivia (L. nubeculosa and L. maya) and Nicaragua (L. fossetti sp. n. and L. cursor). This may be associated with a species' size and/or speed and how quickly it can escape potential predators; slower species requiring smaller diameter trunks in order to disappear to the opposite side quickly. Trunk diameter preferences were found in Ciulfina biseriata Westwood, 1889, an Australian liturgusid bark mantis (O'Hanlon 2011), which could indicate similar preferences within the highly similar Neotropical bark mantises.
Specimens of Liturgusa have been collected by the author and colleagues in wet, low elevation rainforests in Peru, Bolivia, Nicaragua, Costa Rica, French Guiana, and Panama. However, L. maya was readily collected in seasonal deciduous forests in western Nicaragua during the dry season, which is interesting considering that L. maya has been sampled almost entirely within rainforest habitats. Although most species appear to be distributed within rainforests, there does not seem to be a strict adherence to habitat type across all of the species within Liturgusa. The forest types for species of Corticomantis gen. n., Fuga gen. n., and Velox gen. n. is unknown.

Geographic range
With limited previous work on Liturgusa, other than the original species descriptions, the geographic distributions are not well known. A limited number of surveys and taxon lists have included various species of Liturgusa, but it is likely that many of the species identifications within these works are not congruent with the findings of this study. For example, the historical understanding of the distribution of L. cayennensis is far broader (Central America to French Guiana) than the current reconstruction of the species distribution (restricted to region immediately surrounding French Guiana). Therefore, most locality records of species cited within taxon lists have been ignored within this study in favor of establishing species distributions directly from georeferenced specimens examined for this study.
After gathering over 500 specimens for this study the range assessment for each species provides the most accurate measures of distribution to date. All species treated within this study are restricted to the Neoptropical region. The northernmost record is that of L. maya with a specimen collected in Mexico, a little north of Mexico City near euparal or placed in glycerin inside microvials and pinned to the specimen. For species for which males are known, verbal descriptions of the male genital complex are provided. Nomenclature for the male genital complex follows Klass (1997) for the following features that were determined as informative for species diagnosis and description for this revision (see Fig. 51B.1 for labeled structure): apofisis falloid (afa), right dorsal phallomere (fallomero dorsale di destra; fda), main body of ventral left sclerite (L4A), main body of dorsal left sclerite (L4B), apical process (processo apicale; paa), distal process (processo distale; pda), ventral plate (piastra ventrale; pia), ventral process (processo ventrale sclerificato; pva), first sclerite of right phallomere (R1).
Measurements. A total of 335 specimens were measured using a Leica M165C stereo-microscope and an IC80 HD coaxial video camera using the live measurements module of the Leica Application Suite (LAS). Measurements captured in this study are extremely precise and approaching or matching this precision will be necessary to use the data to properly diagnose a specimen. In addition, accurate measurements are critical in the diagnostic key and without them species level identification will be much more difficult. A series of unique measurement metrics are present in the diagnostic key that are capable of separating species or groups of species. The definitions of these metrics are provided within the key itself and usually refer to one of the collected measurements or ratios described below.
All measurements presented in this study are in millimeters. A total of 21 measurement classes were captured including:

1.
Body length = length of body from central ocelli to posterior tip of wing or abdomen (intraspecifically variable measurement, primarily for general size estimation).

2.
Forewing length = from proximal margin of axillary sclerites to distal tip of the discoidal region.

3.
Hindwing length = from proximal margin of axillary sclerites to distal tip of the discoidal region.

4.
Pronotum length = from anterior margin to posterior margin.
Pronotum width = from lateral margins at the widest point, the supra-coxal bulge. 7.
Pronotum narrow width = from lateral margins of the pronotum at narrowest region of metazone. 8.
Head width = from lateral margins of the eyes at widest point. 9.
Head vertex to clypeus = from the vertex of the head at center to the lower margin of the frons and upper margin of clypeus. 10. Frons width = from lateral margins of frons, inferior to the antennal insertions, at the widest point. 11. Frons height = from upper margin abutting central ocellus to lower margin abutting clypeus. 12. Prothoracic femur length = from proximal margin abutting trochanter to distal margin of genicular lobe.
13. Mesothoracic femur length = from most proximal margin abutting trochanter to the distal side of the terminal spine insertion site. 14. Mesothoracic tibia length = from most proximal groove near joint with the femur to the distal side of the terminal spine insertion site. 15. Mesothoracic tarsus length = from proximal joint to the apex of the ungues curve. 16. Metathoracic femur length = from most proximal margin abutting trochanter to the distal side of the terminal spine insertion site. 17. Metathoracic tibia length = from most proximal groove near joint with the femur to the distal side of the terminal spine insertion site. 18. Metathoracic tarsus length = from proximal joint to the apex of the ungues curve. 19. Anteroventral femoral spine count = all inner marginal ridge spines and two proximal near marginal spines, but excluding the genicular spine. 20. Anteroventral tibial spine count = all inner marginal ridge spines, but excluding the distal terminal spur. 21. Posteroventral tibial spine count = all outer marginal ridge spines, but excluding the distal terminal spur.
The measurement of total body length was taken from the central ocellus to tip of posterior margin of abdomen or wing, which produced a variable measure only useful for general assessment of body size rather than species description. Since head position, abdominal expansion, and wing position are all variable, total body length should only be used as a rough measure class to initially discriminate between the small and large species when performing identifications.
A python script was written to provide measurement summaries as well as producing four ratios to provide informative measures of shape characteristics including:

1.
Pronotum elongation measure = prozone length over total length of pronotum (a low measure indicates an elongate metazone).

2.
Pronotum shape measure = pronotum width at the supra-coxal bulge over total length of pronotum (a low measure indicates an elongate pronotum).

3.
Head shape measure = length of vertex of head to lower margin of frons over width of head (a low measure indicates a highly transverse head).

4.
Frons shape measure = height of frons over width of frons (a low measure indicates a highly transverse frons).
For each species and sex, the specimen number (e.g. N=12) of measured individuals is presented prior to the summary measurements, which includes the range and the mean (for quantitative measurement classes) or mode (for meristic measurement classes) for each measurement class (e.g. 29-31 (30)). In cases where more than 15 specimens were present for any species, a subset was taken for measurements that was representative of total geographic range and size variation. Occasionally, certain class measures were not possible to collect (visibility of feature or absence of feature) and are reflected by their absence in the summary or the absence of a range in cases of multiple specimens examined.
The ultimate diagnostic characters to be used in combination include measurements, the structure of male genitalia, and shape of the pronotum. The morphology of the head and prothoracic legs are also helpful in a number of cases. Coloration patterns are used in this study, but these are conservative. If coloration is used in the description, the patterning was consistent across all examined specimens, but coloration not included was typically variable within species. Preservation of specimen, population level variation, and specimen age are all contributors to color alterations and make such characters somewhat untrustworthy. The vast majority of coloration patterning is not discussed as intraspecific variation makes them unreliable for delimiting a species.
Imaging. High resolution images of type and voucher specimens were captured using a Passport Storm© system (Visionary Digital™, 2012), which includes a Stackshot z-stepper, a Canon 5D SLR, macro lenses (50mm, 100mm, and MP-E 65mm), three Speedlight 580EX II flash units, and an associated computer running Canon utility and Adobe Lightroom 3.6 software. The z-stepper was controlled through Zerene Stacker 1.04 and images were processed using the P-Max protocol. All images were captured over an 18% grey card background for white balance standards. Images were processed in Adobe Photoshop CS6 Extended to adjust levels, contrast, exposure, sharpness, and add scale bars (10 mm). Minor adjustments were made using the stamp tool to correct background aberrations and to remove distracting debris. Plates were constructed using Adobe Illustrator CS6.
Illustration. Two dorsal habitus illustrations were produced by sketching on tracing paper before scanning into digital form with a flatbed scanner (Fig. 2). The sketches were imported into Adobe Illustrator to produce a full body fill before using Adobe Photoshop for detail work. Diagrammatic illustrations were produced by collecting reference images of the specimens using both the Leica M165C stereo microscope paired with the IC80 HD camera as well as the Passport Storm, Visionary Digital system. Images were imported into Adobe Illustrator and traced using an Intuos4 drawing tablet. Adobe Illustrator was used for all plate layouts. All illustrations (except the habitus of Liturgusa maya by Julio Rivera, Fig. 12B) were produced by Joshua Maxwell of the Cleveland Institute of Art.
Type designation. Upon creation of the genus Liturgusa by Saussure in 1869, two species, Mantis annulipes Audinet Serville, 1838 and L. cayennensis Saussure, 1869, were included, but neither were designated as the type species for the genus. Further, no type was established until subsequent designation, adherent to Article 69.1 of the Code, by Kirby (1904: 271) of L. cayennensis Saussure, 1869, which was valid under Article 67.2 of the Code as this species was an "originally included nominal species" available for fixation. However, Giglio-Tos (1927: 292) took subsequent action by designation of Mantis annulipes Audinet Serville, 1838, an act not valid according to Article 69.1.2 of the Code, which states that the first designation in a publication (Kirby 1904) is to be accepted. This type discrepancy was first recognized by Rehn (1935: 198) having stated in footnote " Giglio-Tos (Das Tierreich, Lief. 50, p. 292, (1927)), erroneously gives annulipes as the genotype. Kirby's fixation is the first, and, being made on one of the two originally included species, must be followed". Unfortunately, recognition of L. cayennensis Saussure, 1869 as the type for the genus has not been uniform across taxonomic works (e.g. Ehrmann 2002 recognizes M. annulipes andOtte andSpearman 2005 recognizes L. cayennensis).
Redescription of the genus. Body: The overall coloration of all Liturgusa species varies within a mottled or camouflage pattern that incorporates black, brown, pale tan, white or grey, and sometimes shades of green. The mottled patterns can be diffuse or highly contrasting with whitish regions abutting black spots or splotches. All species are dorsoventrally flattened with disproportionately long legs in comparison to body length.
Head: Wider than long with large, rounded eyes projecting outside the profile of the head both laterally and anteriorly (the anterior margin of the eyes anterior to the central surface of the head). Juxta-ocular protuberances present to varying degrees within males, but always well developed in females. The vertex between the parietal sutures is straight, concave or slightly irregular. Frontal suture with a medial carina. Ocelli present in males, but size is variable, protruding on cuticular mounds; reduced in females and laying more flatly on the surface or all three positioned laterally on a continuous curved carina. Central ocellus oriented anteriorly and lateral ocelli oriented outward, perpendicular to the central axis of the head or at most a few degrees off perpendicular. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus convex, lower margin straight, concave, or convex; a transverse ridge medially; lateral margins tapering, widest at the upper margin. Labrum with minimal sculpting and a rounded terminus. Antennae filiform and with rare setae, pale or dark or a combination of both, never banded. Varying levels of black markings across the anterior surface that can include a transverse band or spots on the lower part of the frons, markings around the ocelli and the vertex, and markings on the clypeus, labrum and mandibles. Palpi are usually pale with or without a darkened terminus.
Pronotum: Varying from elongate (pronotum shape measure 0.28) to squat (pronotum shape measure 0.52) with a defined supra-coxal bulge; dorsal surface mostly smooth or at most with disperse tubercles, particularly in the posterior half. Prozone with lateral margins that are parallel, tapering anteriorly or rarely convex. Metazone with lateral margins that are parallel, concave, or tapering posteriorly; the dorsal surface often with laterally symmetrical bulges in the middle, which can push lateral margins outward. Coloration highly variable with pale and black markings. Supra-coxal sulcus strongly defined; posterior margin straight or medially emarginate.
Prothoracic Legs: Femoral spine count of male and female: anteroventral 12-17, posteroventral 4, discoidal 4. Femur robust with a straight or concave dorsal margin; anteroventral and posteroventral (internal and external, respectively) spines well developed; line of small tubercles running medially of the posteroventral spines. A continuous carina running from distal terminus of femur along dorsal margin to the base, circling the external surface of the proximal end and running along the ventral margin at the base of the posteroventral spines. Pale to dark banding on posterior (external) surface of femur; anterior (internal) surface pale with varying patterns of black markings. Posterior surface of femur smooth or with few tubercles. Well-developed femoral pit to accommodate terminal posteroventral tibial spine, positioned medial to the proximal two posteroventral spines; pit is colored black, brown or pale. Prothoracic tibial spine count of male and female: anteroventral 7-11, posteroventral 7. Prothoracic tibial spines robust; the posteroventral spines with the first smallest, the second and/or third longer, the third or fourth through sixth of similar length; the anteroventral spines longest at distal end and shortening proximally, but the seventh and eighth spines from the distal terminal spine longer than adjacent spines. Tarsus normal for Mantodea, but banded with pale and dark coloration. Prothoracic coxae smooth with no or a few very minor tubercles or setae along dorsal margin; black markings vary across species on the anterior, posterior, and ventral surfaces.
Meso-and Metathoracic Legs: Long and slender with pale to dark banding on the femur and tibia; dorsal surface of femora smooth. Femora with ventral (posterior) carina, some species being more pronounced than others; dorsal (anterior) carina noticeable in some species (Cayennensis Group in particular). Tibia long and rounded with well developed terminal spurs. Mesothoracic tarsi with first segment shorter, equal to or longer than remaining segments combined. Metathoracic tarsi with first segment always at least slightly longer than remaining segments combined, can be much longer.
Wings: Wings developed in males and females. Forewing mottled with brown, black, white, and green coloration; the costal region narrow relative to the wing length, the width between 2-5% the length, often with light to dark irregular banding; veins often marked with irregular sections of pale color. The forewings in many species may be colored asymmetrically, one being mottled as described above while the other is either dark rust or blackened with the mottled pattern still slightly visible (darker wing typically folded under the mottled wing). Hindwings hyaline, smoky opaque, and/ or with rusty, yellow, or orange coloration; the terminus of the discoidal region either projecting beyond or within the profile of the distal margin of anal region.
Abdomen: Males and females with varying degrees of gradual widening from first segment until the beginning of the distal third (segments 5-7) at which point the lateral margins narrow to the terminus, the middle third being the broadest region. Some slender species with very slight widening, exhibiting near parallel margins before an abrupt narrowing as described above. Some species with pointed posterolateral tergal projections in the distal half of the abdomen of males and/or females, but other species with unmodified tergites. Cerci cylindrical, long and setose, tapering to a point. Supra-anal plate long or transverse, always with a rounded terminus of varying degrees. Subgenital plate of male with rounded, slightly irregular terminus; without styli.
Male Genital Complex: The distal end of main body of ventral left sclerite (L4A) is either smooth and rounded or with a distal process (pda) of varying size and shape. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) well sclerotized with a highly variable terminus; the apical process (paa) cylindrical and curved, terminating in a rounded or blunt end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus and is mostly membranous with disperse setae of varying robustness; the ventral plate (pia) strongly sclerotized with strongly defined grooves, slight grooves, or smooth; the ventral process (pva) strongly sclerotized and curved and/or tapering distally.
Ootheca (Figs 1D,. For known species, the ootheca is attached to solid substrate, usually tree bark. The main body is spherical with a tapering hollow tube originating from a dorso-medial position at about 45 degrees from a lateral perspective relative to attachment surface. Eggs positioned in a linear row medially in line with the tubicular process within an air-filled space. The size and volume of the air-filled space of the spherical body appears to vary across species (a considerably larger ootheca was examined for an unknown species in the Natural History Museum, London). Upon hatching, nymphs emerge through the tube to the outside. The number of eggs is unknown across the species.
Description. Male. (Fig. 3A  7.14-8.20 (7.43); mesothoracic tibia length 5.34-6.38 (5.70); mesothoracic tarsus length 4.33-5.71 (5.13); metathoracic femur length 6.75-8.36 (7.43); metathoracic tibia length 6.05-8.51 (7.49); metathoracic tarsus length 6.24-8.17 (7.10); pronotal elongation measure 0.30-0.34 (0.32); pronotal shape measure 0.44-0.46 (0.45); head shape measure 0.37-0.40 (0.39); frons shape measure 0.37-0.44 (0.40); anteroventral femoral spine count 12-16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7. Head ( Fig. 40A): Transverse, the juxta-ocular protuberances small, but pronounced, the apex in the lateral half; the vertex is straight, even with the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, most pronounced medially, the region just ventral depressed. Ocelli small and protruding on small cuticular mounds, but the region between all three slightly raised; the lateral ocelli oriented outward. The carina on the frons pronounced, the medial region just ventral to the carina depressed. Clypeus transverse, the upper margin convex, the lower margin slightly concave; the central carina strongly pronounced and straight. Antennae pale at the base, the flagellum fading to dark brown just slightly distal to the base. Black band extending straight over the medial carina of the frontal suture, the medial portion of the carina pale; a branch of the black band extends ventro-laterally between the eye and antennal insertion at a forty five degree angle; a branch of the black band extends dorsal-medially from the lateral region of the main transverse band and terminating near the ventral terminus of the parietal suture. Lower region of frons darkened; the clypeus is pale; the mandibles and labrum mostly pale, but with some brown marks; the vertex and juxta-ocular protuberances pale and brown; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47A): Short and squat with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone squat with margins gradually tapering anteriorly to a rounded anterior margin; the lateral margins smooth. Metazone with concave lateral margins, tapering posteriorly until about two thirds distance from supracoxal bulge then widening gradually to the posterior margin; margins smooth; posterior margin with a slight medial emargination; the dorsal surface of the posterior third of the metazone depressed; two small and elongate bulges present on the dorsal surface near the posterior margin and positioned laterally. Pale with strong black marks across the surface, two prominent black marks laterally just posterior to the supra-coxal sulcus.
Prothoracic Legs: Femur squat and robust with a near straight dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a very thin black band running medially from the base to terminus, a black mark dorsal to the band at the midpoint, and a dark mark dorsal to the band near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and exactly between the first two proximal posteroventral spines, but slightly distal to the most distal discoidal spine; pit is pigmented darkly. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are 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 pronounced. Mesotarsi with first segment as long as the remaining segments combined.
Wings: Forewings mottled with brown, pale and greenish coloration; the costal region without strongly defined banding, green and brown proximally with some low contrasting bands developing distally; veins are brown with cells being dark brown or light brown; two pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein; a large pale spot 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 opaque brown, the discoidal region more pale proximally; the costal region light brown proximally, darkening distally; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing slightly elongate.
Abdomen: Broad, widening until the fifth tergite before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, a rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51A.1-A.2): The main body of ventral left sclerite (L4A) with rounded terminus, but with a short, laterally positioned distal process (pda) that is rounded and sometimes projecting at an angle towards the medial axis of the L4A that can create a strongly angled transition from the terminal margin of the L4A to the medial margin of the pda; sometimes a depression on the lateral half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) short, broad and tapering to a point and heavily sclerotized, often curved; the apical process (paa) short, cylindrical and curved, the terminus a 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 with a rough surface, but mostly lacking strongly defined grooves though sometimes present; the ventral process (pva) smooth and tapering to a point distally, one edge straight and the other convex, tooth-like in appearance.
Head (Fig. 40B): As long as wide, the juxta-ocular protuberances very large, the apex in the middle; the vertex is straight, well above the dorsal margin of the eyes.
Pronotum (Fig. 47B): Dorsal surface entirely smooth except from a rough texture in the posterior quarter, no defined tubercles. Metazone with very few, small tubercles centrally located.
Prothoracic Legs: A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and exactly between the first two proximal posteroventral spines, and in line with the most distal discoidal spine. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are slightly longer (the second much longer than the third). Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe.
Wings: Forewings with a widened costal region. Forewings extending to the tip of the abdomen or shorter (depends largely on preservation of specimen). Hindwings pale along the anterior margin and the distal terminus of the discoidal region; the discoidal region broadened.
Abdomen: Broad, widening from first segment until the beginning of the distal half (segment 5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supraanal plate slightly transverse, and evenly rounded lobe.  : Westwood 1889: 5, 51;Kirby 1904: 271;Caudell 1918: 5;Rehn 1935: 199;Rehn 1954: 179, pl Taxonomic history. Described in 1889 by Gerstaecker, the species has been largely ignored other than inclusion in taxonomic lists and some regional studies. Giglio-Tos (1927) considered L. lichenalis as the synonym of F. annulipes, but this treatment is erroneous as it is apparent that no previous work had an accurate concept of F. annulipes. Rehn (1935) corrected this erroneous synonym by comparing the types of both species.

Liturgusa lichenalis
Diagnosis. Almost identical to L. cayennensis, but primarily distributed in the western Amazon basin rather than in the Guyanas in the northeastern coastal region of South America. In addition, the abdomens of males and females are broader than L. cayennensis and have posterolateral tergal projections in the distal half. Another distinguishing external character for L. lichenalis comparison to L. cayennensis is that the first segment of the mesotarsi is obviously shorter than the remaining segments combined while this segment is the same length as the remaining segments combined in L. cayennensis.
Head ( Fig. 40C): Transverse, the juxta-ocular protuberances small, but pronounced, the apex slightly in the lateral half; the vertex is straight, even with the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, most pronounced medially, the region just ventral depressed. Ocelli small and protruding on small cuticular mounds, but the region between all three slightly raised; the lateral ocelli oriented outward. The carina on the frons present, but not highly pronounced, the medial region just ventral to the carina depressed. Clypeus transverse, the upper margin convex, the lower margin slightly concave; the central carina strongly pronounced and straight. Antennae pale at the base, the flagellum fading gradually to dark brown around a third of the way from the base. Black band extending straight over the medial carina of the frontal suture, the medial portion of the carina pale; a branch of the black band extends ventro-laterally between the eye and antennal insertion at a forty five degree angle; area around parietal sutures pale or dark brown. Lower region of frons darkened; the clypeus is pale; the mandibles and labrum mostly pale, but with some brown marks; the vertex and juxta-ocular protuberances mostly pale; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47C): Short and squat with a poorly defined supra-coxal bulge; dorsal surface entirely smooth or with few blunt tubercles in the posterior half. Prozone squat with near parallel margins before tapering anteriorly to a rounded anterior margin; the margins smooth. Metazone with barely concave lateral margins, mostly tapering consistently from the supra-coxal bulge posteriorly until a very slight widening in the posterior quarter leading to the posterior margin; margins smooth; posterior margin with a slight medial emargination; the dorsal surface of the posterior half of the metazone depressed; two small and elongate bulges barely present on the dorsal surface near the posterior margin and positioned laterally, sometimes absent. Pale with strong black marks across the surface, two prominent black marks laterally just posterior to the supra-coxal sulcus.
Prothoracic Legs: Femur squat and robust with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a very thin black band running medially from the base to terminus, a small black mark dorsal to the band at the midpoint, and a dark mark dorsal to the band near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned slightly distal to the first proximal posteroventral spine and in line with the most distal discoidal spine; pit is pigmented darkly or pale. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are slightly longer. Prothoracic coxae smooth, the anterior surface pale.
Wings: Forewings mottled with brown, pale and greenish coloration; the costal region without strongly defined banding, green and brown proximally with some low contrasting bands developing distally; veins not contrasting from surrounding coloration; two pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein; a large pale spot is positioned centrally. Forewings often, but not always asymmetrically colored; one being mottled as described the other is darkened significantly with a rust tone, the mottled pattern still visible; extending just beyond or as long as the abdomen. Hindwings with discoidal region opaque and colored brown or rust; the anal region smoky and translucent; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing slightly elongate.
Abdomen: Broad, widening until the fifth tergite before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites with small posterolateral tergal projections beginning on the sixth segment. Supra-anal plate slightly transverse, margins tapering gradually to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51B.1-B.4): The main body of ventral left sclerite (L4A) with rounded terminus, but with a short, laterally positioned distal process (pda) that is rounded and sometimes, but not always, projecting at an angle towards the medial axis of the L4A that can create a strongly angled transition from the terminal margin of the L4A to the medial margin of the pda; sometimes a depression on the lateral half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) very short, broad, tapered to the point and heavily sclerotized, not curved; the apical process (paa) short, cylindrical and curved, the terminus with a rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a broad, rounded, membranous terminus, the lateral margin often folded over; the ventral plate (pia) long, broadened proximally and with strongly defined grooves; the ventral process (pva) smooth and tapering to a point distally, one edge straight and the other convex, tooth-like in appearance, but the proximal half c-shaped.
Head (Fig. 40D): As long as wide, the juxta-ocular protuberances very large, the apex in the middle; the vertex is slightly concave, above the dorsal margin of the eyes. Ocelli small and protruding on a continuous carina connecting the three and extending slightly laterally; region between all three raised slightly. Black marking present over the medial carina of the frontal suture; the vertex and juxta-ocular protuberances with dark markings.
Pronotum ( Fig. 47D): Metazone with barely concave lateral margins, tapering rapidly in the first third from the supra-coxal bulge, the middle third near parallel before widening to the posterior margin; margins smooth or with few small tubercles; posterior margin with a medial emargination; few small tubercles present.
Prothoracic Legs: Anterior (internal) surface of femur with a very thin black band running medially from the base to terminus that is often interrupted, a small black mark dorsal to the band at the midpoint, and a dark mark dorsal to the band near the femoral brush.
Meso-and Metathoracic Legs: As described for males. Wings: Forewings mottled with contrasting brown, pale and greenish coloration; veins pale brown. Forewings often, but not always asymmetrically colored; one being mottled as described the other is darkened with a rust or black tone, the mottled pattern still visible; extending to the terminus of the abdomen. Hindwings with discoidal region opaque and colored brown, rust or dark yellow; the anal region smoky and translucent.
Abdomen: Very broad, widening until the fifth tergite before a gradual posterior narrowing, the abdomen almost circular; a smooth, brown and black colored dorsal surface. Tergites with large posterolateral tergal projections beginning on the fourth segment. Supraanal plate slightly transverse, margins tapering gradually to a rounded terminus. Taxonomic history. Described in 1939 by Marcello La Greca, L. guyanensis was only subsequently treated in taxon lists and has received no revisionary attention. La Greca only included the type locality, but no repository for the type. It is presumed that the specimen remains in an Italian collection. In addition, the species was cited in subsequent works with the repository listed as the Natural History Museum Zoological Section 'La Specola', University of Florence. Therefore, Luca Bartolozzi was contact in hopes of locating the specimen. After an extensive search it was not located in Florence. Luca Bartolozzi contacted museums in Genoa, Milan and Rome looking for the specimen, but nothing was located in these collections. After consulting with Luca Picciau, of the Museo Regionale di Scienze Naturali, Torino, in search of the type specimen of L. charpentieri, described and deposited there by Giglio-Tos (1927), it was learned the specimen was loaned to La Greca, but after his death his entire collection, presumably including loans, was sold to the Museo Civico di Storia Naturale, Milan. Chasing this lead, Fabrizio Rigato and Michele Zilioli, both of the Museo Civico di Storia Naturale, Milan, were contacted, but neither the type of L. charpentieri nor L. guyanensis were located in the collection. In addition, the eminent mantodean systematist, Prof. Francesco Lombardo of the Università degli Studi di Catania, was consulted throughout this search, but he as well was not able to locate the type. Therefore, in satisfying Article 75.3 of the International Code of Zoological Nomenclature, a specimen has been located that matches the original description and illustrations provided by La Greca (1939) and represents this distinct species within Liturgusa. Although La Greca's type was female and the designated neotype is male, a female specimen has also been located and is treated herein, which demonstrates a match with the original name-bearing type. A male is designated as the neotype herein as a measure of consistency across the entire genus, which now includes holotype descriptions of male genital structures. Finally, the neotype is not from the original type locality, but from just south within Guyana and still on the eastern side of the Guyana Shield, within the same elevational range, forest type, and climate conditions.

Liturgusa guyanensis
Diagnosis. Similar to L. cayennensis and living in sympatry with at least some populations of the species, L. guyanensis have a far more sculpted pronotum, a unique vertex of the head (straight with lateral depressions near the parietal sutures), and evenly mottled forewings (without the large pale or whitish regions that are obviously present on L. cayennensis and L. lichenalis).
Head ( Fig. 40E): Slightly transverse, the juxta-ocular protuberances moderately pronounced, the apex in the lateral half; the vertex is convex, slightly higher than the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, the region just ventral depressed for most of the length. Ocelli small and protruding prominently on small cuticular mounds; the lateral ocelli oriented outward. The carina on the frons pronounced, the medial region just ventral to the carina depressed and sloped ventrally. Clypeus transverse, the upper margin convex, the lower margin straight; the central carina strongly pronounced and straight. Antennae pale at the base, the flagellum absent on specimen. The vertex and juxtaocular protuberances pale and with brown speckled markings; the area immediately adjacent to lateral ocelli black; the ventrolateral margins of the frons with a dark brown margin. Palpi are pale.
Pronotum (Fig. 47E): Short and squat with a defined supra-coxal bulge; dorsal surface with disperse, small tubercles. Prozone squat with convex margins widening anterior to supra-coxal bulge before narrowing to the anterior margin; the margins smooth. Metazone with sweeping concave lateral margins, the nadir at the three quarter point from the supra-coxal bulge, then widening slightly to the posterior margin; margins with small, disperse tubercles; posterior margin with a slight medial emargination, almost straight; the dorsal surface of the posterior third of the metazone depressed; tubercles more common in the posterior half. Pale with strong black marks across the surface, two prominent black marks laterally just anterior to the supra-coxal sulcus in the prozone.
Prothoracic Legs: Femur squat and robust with a near straight dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a very thin black band running medially from the base to terminus, a black mark dorsal to the band at the midpoint and some thickening of the line near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned just distal to the first most proximal posteroventral spine and in line with the most distal discoidal spine; pit is pigmented darkly. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are slightly longer. Prothoracic coxae smooth, the anterior surface with a very small, black mark in the proximal half and adjacent to the medial line.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina pronounced. Mesotarsi with first segment slightly longer than the remaining segments combined.
Wings: Forewings evenly mottled with dark and light brown coloration; the costal region without strongly defined banding, mostly matching the color patterns of the discoidal region; veins are pale, contrasting with the cell colors. Forewings asymmetrically colored; one being mottled as described the other is darkened significantly with a rust tone, the mottled pattern still visible; extending just beyond the abdomen. The discoidal region of the hindwings opaque, a pale rust color proximally that fades to a dark, rusty opaque color in the distal half; the anal region of the hindwing smoky and translucent; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Broad, widening until the fifth tergite before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites without posterolateral tergal projections. Supra-anal plate transverse, a broadly rounded terminus with a medial emargination. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51C.1): The main body of ventral left sclerite (L4A) with a convex terminal margin that tapers to a medially positioned, blunt point that is well sclerotized, but lacking a distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate, slender and curved, terminating in a blunt, but narrow point; the apical process (paa) short, cylindrical and curved, the terminus with a 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 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.
Head ( Fig. 40F): Approximately as broad as wide, the juxta-ocular protuberances very large, the apex in the middle; the vertex is straight, but with two depressions just medial to the parietal sutures, higher than the dorsal margin of the eyes. Ocelli very small and protruding prominently on small cuticular mounds, region between ocelli, ventral to the frontal suture and dorsal of the frons is depressed. Clypeus transverse, the upper margin convex, the lower margin rounded. Antennae pale at the base, the flagellum fading to brown gradually.
Pronotum (Fig. 47F): Prozone squat and broad with convex margins widening prominently anterior to supra-coxal bulge before narrowing quickly to the anterior margin; the margins with small tubercles. Metazone with concave lateral margins, the nadir at the three quarter point from the supra-coxal bulge, but a slight bulge in the posterior half present that pushes the margins laterally and interrupts the continuity of the concave margins; margins with small, numerous tubercles.
Prothoracic Legs: Femur with anterior (internal) surface with a black band running medially from the base to terminus, a black mark dorsal to the band at the midpoint and some thickening of the line near the femoral brush, but the overall band may be interrupted medially. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are longer.
Meso-and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Costal region of forewing widened. Forewings symmetrically colored; extending the length of the abdomen. The terminus of the discoidal region of the hindwing projecting slightly beyond the distal margin of anal region, the wing appearing slightly elongate.
Abdomen: Broad, widening from first segment until the beginning of the distal half (segment 5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supraanal plate slightly transverse, and evenly rounded lobe. Material examined. Liturgusa neblina sp. n.

Diagnosis.
Most similar in general appearance to L. cayennensis and L. lichenalis. The type locality overlaps with the range of L. lichenalis, but L. neblina is distinct in a few ways. First the hindwings are yellow-orange in the discoidal region with either a rapid or slow fade to a hyaline or translucent brown in the anal region. The pronotum is short and squat as are the other species within the Cayennensis Group, but the metazone is more constricted. The forewings are evenly mottled with green and brown with regular pale marks in the distal half.
Head ( Fig. 40G): As long as broad, the juxta-ocular protuberances very large, the apex in the middle; the vertex is slightly concave, above the dorsal margin of the eyes. Frontal suture with a very faint medial carina forming a continuous arc, may be seen primarily because of pale coloration compared to the dark markings above and below. The region ventral to the frontal suture depressed, gradually sloping higher to the central ocellus. Ocelli small and protruding on a carina that connects all three and extends laterally a short distance; the lateral ocelli oriented outward; area just ventral to central ocellus depressed (upper region of frons). The carina of the frons pronounced, the medial region just ventral to the carina depressed, sloped ventrally. Clypeus slightly transverse, the upper margin slightly convex, the lower margin concave; the central carina strongly pronounced and straight. Antennae is pale at the base, the flagellum absent from specimen. Curved, black band extending over the frontal suture, the carina pale; projections from the band extend dorsally from the middle, ventrally toward the central ocellus, and surrounding the lateral ocelli and extending laterally above the antennal insertion. Lower region of frons darkened; the clypeus is pale; the mandibles mostly pale, but with some brown marks distally; the labrum pale but with two laterally position black marks near the dorsal margin; the vertex and juxta-ocular protuberances mottled with pale and brown. Palpi are pale.
Pronotum (Fig. 47G): Short and squat with a moderately defined supra-coxal bulge; dorsal surface with a few blunt tubercles in the posterior half. Prozone squat with tapering margins anteriorly to a rounded margin; the margins smooth. Metazone with posteriorly tapering margins until the posterior half where the margins are parallel before widening again prior to the posterior margin; posterior two thirds of the margins with small tubercles; posterior margin with a slight medial emargination; the dorsal surface of the posterior half of the metazone depressed. Pale with strong black marks across the surface, two prominent black marks laterally just anterior to the supra-coxal sulcus. Prothoracic Legs: Femur squat and robust with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface mostly pale, but with a faint dark mark medially and the distal third with a black band running along the medial line with an expanded region near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to the two most proximal posteroventral spines and in line with the most distal discoidal spine; pit is pigmented brown. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are longer, but the second is very long. Prothoracic coxae smooth, the anterior surface pale.
Wings: Forewings evenly mottled with brown and green, without large contrasting regions of pale marks; the costal region without strongly defined banding, green and brown mottling; veins green and contrasting from surrounding coloration; two pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein. Forewings asymmetrically colored, one being mottled as described the other is darkened significantly with a rust tone, the mottled pattern still visible; extending slightly beyond the abdomen. Hindwings with discoidal region opaque and colored yellow or orange, darkening distally; the anal region with a very narrow anterior margin colored as in the discoidal region, hyaline otherwise; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing slightly elongate, but still broad.
Abdomen: Broad, widening until the fifth tergite before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, margins tapering gradually to a rounded terminus.
Etymology. A noun in apposition, L. neblina is from the Cerro de la Neblina, the geological formation giving rise to the Pico de Neblina, a tepui located in southern Venezuela near the border with Brazil. The only known female was collected at the basecamp for Pico de Neblina. Natural history. Only three individuals (one escaped) were observed in two locations in the Loreto Province, Peru. Two females, both were found less than one meter from the ground on small to medium diameter, smooth bark trees with patches of moss. The female that escaped ran into the herbaceous vegetation surrounding the tree at the base and disappeared. The collected female was easily corralled low on the tree. The male was found living in sympatry with L. algorei, L. krattorum, and L. maya at the Madre Selva Biological Research Station in the Loreto Province, Peru.
Head ( Fig. 41A): Transverse, the juxta-ocular protuberances small, the apex in the lateral half; the vertex is nearly straight, but with two slight bulges just medial to each parietal suture; vertex just below the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc, the region ventral to the carina, particularly in the middle, is depressed forming a deep groove. Ocelli small and protruding slightly on small cuticular mounds; the central ocellus slightly larger than lateral; than lateral ocelli oriented outward. The carina on the frons not pronounced, but present. Clypeus transverse, the upper margin slightly convex, the lower margin straight or slightly concave; the central, transverse carina pronounced, straight. Antennae scape and pedicel pale, the flagellum light brown or black. Area around frontal suture, vertex and the juxta-ocular protuberances brown with black or darker markings, no distinct transverse band. Lower region of frons dark brown, a small pale region along ventral margin; clypeus pale; the mandibles and labrum pale with darker brown markings. Palpi are pale.
Pronotum (Fig. 47H): A little less than three times as long as wide with a moderately defined supra-coxal bulge; dorsal surface mostly smooth, but sometimes with rough patches or small tubercles in the posterior third. Prozone square with slightly convex margins that gradually taper to an evenly rounded anterior margin; margins smooth or with blunt tubercles. Metazone with concave lateral margins; margins smooth or with small tubercles; the dorsal surface of the posterior third of the metazone depressed. Mostly pale or brown with darker brown or black markings across the surface.
Prothoracic Legs: Femur normal with a concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface darkly pigmented, only two pale marks along the dorsal margin; the ventral surface darkly pigmented with brown or black. Posterior surface of femur with few tubercles. A relatively large and shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and just distal to the proximal most posteroventral spine, in line with the most distal discoidal spine; pit is pigmented dark brown. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating just proximal of 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 longer. Prothoracic coxae smooth, the anterior surface with a small, black mark medially in the proximal half as well as a 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 shorter or as long as the remaining segments combined.
Wings: Forewings mottled with pale, green, and brown coloration; the costal region mostly pale with a few dark marks, no regular banding; vein coloration is pale, but sections match darker surroundings; discoidal region darker proximally, the distal half more pale in overall color dominance. Forewings not asymmetrically colored; extending just beyond the terminus of the abdomen. Hindwings with opaque brown or black discoidal region, darker proximally; the anal region smoky black and translucent; the terminus of the discoidal region projecting far beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Slightly widened, the fifth or sixth 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 transverse, evenly tapered terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51D.1): The main body of ventral left sclerite (L4A) asymmetrical, the left margin rounded, leading to a short, blunt nub projecting laterally; the right margin of L4A straight; lacking a distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate and robust, one margin being concave and connecting to a prominent subprocess of the L4B, the entire structure cshaped; the apical process (paa) elongate and thick, the terminus evenly rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a broadly rounded, membranous terminus; the ventral plate (pia) short and broad with a rough surface and poorly defined grooves; the ventral process (pva) small and c-shaped, the distal terminus tapered to a point.
Head ( Fig. 41B): Transverse, the juxta-ocular protuberances large, the apex in the lateral half; the vertex is concave; vertex just above the dorsal margin of the eyes. The central ocellus the same size as the lateral ocelli. The carina on the frons pronounced.
Pronotum (Fig. 47I): As described for males. Prothoracic Legs: Anterior (internal) surface of femur darkly pigmented, pale regions along the ventral and dorsal margins; the ventral surface pale, but with two darkly pigmented medial spots, one in line with the second most proximal posteroventral spine and the other just proximal to the most distal posteroventral spine. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating well proximal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) and third the same size, the second significantly longer, the fourth through sixth the shortest and of similar length to each other.
Meso-and Metathoracic Legs: As described for males. Wings: The costal region of forewing mottled with pale and dark colors, irregular banding present; mostly pale with a few dark marks, no regular banding; vein coloration matching surroundings; discoidal region darker proximally, a central pale mark, then dark, turning pale in the distal quarter. Forewings not asymmetrically colored; extending to around the terminus of the abdomen.
Abdomen: Widened, the fourth tergite the widest region before a gradual posterior narrowing. Seventh tergites with small posterolateral tergal projections. Supra-anal plate slightly transverse, a broadly rounded terminus.
Etymology. A noun in the genitive case, L. bororum is named for the Bora people, a marginalized tribe of people native to parts of the Amazon basin in northern Peru, Colombia and Brazil. Their current population is estimated at 2,000 and after a devastating period during the rubber boom, they remain with no indigenous reserves. Diagnosis. The smallest Liturgusa species, L. cura is most similar to L. bororum and L. manausensis, but is located only in northern Venezuela, a unique distribution. Also, L. cura has rounded posterolateral margins of the metazone, distinct from L. manausensis, as well as a mostly pale ventral prothoracic femoral surface (male has a medial brown marking) that is distinguished from L. bororum.
Head ( Fig. 41C): Transverse, the juxta-ocular protuberances small, the apex in the lateral half; the vertex is straight; vertex just below the dorsal margin of the eyes.
Frontal suture with a slight medial carina forming a continuous arc, the region ventral and dorsal to the suture not depressed. Ocelli small and protruding slightly on small cuticular mounds; the central ocellus slightly larger than the lateral ocelli; the lateral ocelli oriented outward. The carina on the frons not pronounced. Clypeus transverse, the upper margin slightly convex, the lower margin straight or slightly concave; the central, transverse carina not very pronounced, straight. Antennae scape and pedicel pale, the flagellum light brown. Area around frontal suture, vertex and the juxta-ocular protuberances brown with black or darker markings, no distinct transverse band. Lower region of frons dark brown, a small pale region along ventral margin; clypeus pale; the mandibles and labrum pale with darker brown markings. Palpi are pale.
Pronotum ( Fig. 47J): Slightly less than three times as long as wide with a moderately defined supra-coxal bulge; dorsal surface mostly smooth, but with tubercles in the posterolateral corners of the metazone. Prozone a little broader than long, the lateral margins nearly parallel, tapering toward a broad and rounded anterior margin; margins smooth or with very small tubercles. Metazone with shallow concave lateral margins; posterior margin with an emargination; margins with small, disperse tubercles; the dorsal surface of the posterior third of the metazone slightly depressed. Mostly pale or brown with darker brown or black markings, two prominent black marks positioned anterolaterally in the metazone.
Prothoracic Legs: Femur normal with a concave dorsal margin; defined pale to dark banding on posterior (external) surface; anterior (internal) surface with an irregular medial band that is interrupted with pale areas; the ventral surface mostly pale, but with a dark brown mark medially between the second and third posteroventral spines. Posterior surface of femur with few tubercles. A relatively large and shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and just distal to the proximal most posteroventral spine, in line with the most distal discoidal spine; pit is pigmented black. Posterior prothoracic femoral genicular spine a little smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) of similar size to the third through sixth, the second longer. Prothoracic coxae smooth, the anterior surface with a small, black mark medially in the proximal half as well as a 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 shorter than the remaining segments combined.
Wings: Forewings mottled with pale, green, and brown coloration; the costal region with regular banding with alternating pale and dark brown, less defined proximally; vein coloration is pale and brown depending on surrounding coloration; discoidal region with a large centrally located pale marking. Forewings asymmetrically colored, one slightly darkened; extending to approximately the terminus of the abdomen. Hindwings with opaque black discoidal region, darker in the anterior half; the anal region smoky black and translucent; the terminus of the discoidal region projecting far beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Slightly widened, the fifth 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, an evenly rounded lobe. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51E.1): The main body of ventral left sclerite (L4A) slightly elongate, the left margin with a large depression near the terminus, the left side with an elongate depression, lacking a distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) short, tapering to a sharp point; the apical process (paa) elongate, the terminus evenly rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus; the ventral plate (pia) small and narrow, not expanded proximally, with a smooth surface; the ventral process (pva) small and c-shaped, with irregular margins.
Head ( Fig. 41D): Transverse, the juxta-ocular protuberances large, the apex just lateral to the middle; the vertex is convex; vertex just above the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc, the region ventral and dorsal to the suture not depressed. Entire vertex above the frontal suture projecting anteriorly, more pronounced than the carina of the frontal suture; two depressions present at the ventral terminus of the parietal suture. Ocelli small and protruding slightly on an elevated carina connecting all three. Clypeus transverse, the upper margin convex, the lower margin concave; the central, transverse carina pronounced, straight. Antennae scape and pedicel pale, the flagellum pale. Area around frontal suture, vertex and the juxta-ocular protuberances mostly pale with brown and black speckling and larger markings, no distinct transverse band. Frons, clypeus, and labrum pale with some brown speckling.
Pronotum (Fig. 47K): As described for males. Prothoracic Legs: Anterior (internal) surface with an irregular medial band that is interrupted with pale areas; the ventral surface pale. A relatively large and shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and just distal to the proximal most posteroventral spine, in line with the most distal discoidal spine; the lateral margin of the pit extends to the lateral margin of the femur, therefore the margin is depressed between the two posteroventral spines; pit is pigmented black. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are longer. Prothoracic coxae smooth; the anterior surface with a small black band medially in the proximal half as well as a small black spot medially towards the distal terminus.
Meso-and Metathoracic Legs: As described for males.
Wings: The terminus of the discoidal region of the hindwing projecting beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Widened, the fifth 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 longer than broad, an evenly rounded lobe.
Etymology. A noun in apposition, L. cura is named from the Villa de Cura in northern Venezuela where the species was collected. Diagnosis. A medium size species with a moderately elongate pronotum, L. fossetti is most similar to L. maya, which is also distributed in Central America. However, L. fossetti can easily be distinguished from L. maya and other Central American species by the yellow coloration on the hindwings.
Head ( Fig. 41E): Transverse, the juxta-ocular protuberances small, 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 medial carina forming a continuous arc, the region just ventral to the carina depressed and the region just dorsal to the carina slightly depressed just lateral to the midline. Ocelli small, the central slightly enlarged, 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 convex, the lower margin slightly concave; the central, transverse carina pronounced and curved. Antennae scape pale, pedicel partly dark brown or black, the flagellum dark brown or black just slightly distal to the base. Vertex and juxta-ocular protuberances mostly dark brown with black marks and black speckling; two prominent pale marks positioned just lateral to the lateral ocelli. Lower region of frons darkly pigmented; the clypeus, labrum, and mandibles pale; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47L): Slightly less than three times long as wide with a moderately defined supra-coxal bulge; dorsal surface smooth, but with a few small tubercles in the posterolateral corners of the metazone. Prozone slightly longer than broad with slightly convex margins that gradually taper to an evenly rounded anterior margin; margins smooth or with very few blunt tubercles. Metazone with shallow concave lateral margins without interruptions or bulges, the medial region near parallel for a short distance; 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 markings across the surface, black marks laterally 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 thin black band running medially from the base to terminus that may be interrupted; the ventral surface pale. Posterior surface of femur with few tubercles. A femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and just distal to the first most proximal posteroventral spine, distal to the most distal discoidal spine; pit is pigmented black. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, 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 longer. Prothoracic coxae smooth, the anterior surface with a black band 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 shorter than the remaining segments combined.
Wings: Forewings mottled with brown, pale and greenish coloration; the costal region with defined banding distally, the proximal region mostly brown; vein coloration across discoidal region pale, not matching surrounding coloration; a pale spot positioned in the proximal quarter of the discoidal region just posterior to the first radial vein; a large pale area is positioned centrally; brown coloration dominant across the discoidal region within cells, the veins pale and appearing like a net-like pattern on the brown background. Forewings often, but not always asymmetrically colored; one being mottled as described, the other is darkened significantly with a rust tone, the mottled pattern still visible; extending just beyond or as long as the abdomen. Hindwings with an opaque yellow coloration in the proximal three quarters, yellow color extending into the anterior area of the anal region, the rest is smoky and translucent; distal quarter of the discoidal region opaque black; 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 transverse, an evenly rounded terminus with a medial emargination. Subgenital plate irregularly rounded and without styli.
Head (Fig. 41F): Slightly transverse, the juxta-ocular protuberances large, the apex in the middle; the vertex is straight, higher than the dorsal margin of the eyes. Antennae scape pale, pedicel dark brown or black, the flagellum dark brown or black just slightly distal to the base. Black band extending straight over the medial carina of the frontal suture, the carina pale; black markings extend ventrally and dorsally from black band. Lower region of frons with dark pigmentation; dorsolateral corners of the clypeus darkly pigmented, the brown pigment extending along the ventral margin of the central carina; the mandibles and labrum with pale and brown markings; the vertex and juxta-ocular protuberances pale with brown speckles; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47M): Dorsal surface smooth, but with tubercles in the posterior half of the metazone. Prozone longer than broad with anteriorly tapering margins.
Prothoracic Legs: Femur robust with a nearly straight dorsal margin; anterior (internal) surface with a degraded (pale interruptions) black band running medially from the base to terminus. A deep femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the two most proximal posteroventral spines, slightly distal to the most distal discoidal spine. Posterior prothoracic femoral genicular spine half the length of posteroventral spines, 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 much longer.
Meso-and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings mottled with brown, pale and greenish coloration; the costal region without defined banding, mostly brown and pale mottled; vein coloration across discoidal region pale, not matching surrounding coloration; a pale spot positioned in the proximal quarter of the discoidal region just posterior to the first radial vein; a large pale area is positioned centrally; the distal half with numerous large pale spots, the background color is dark brown; costal region widened. Forewings not asymmetrically colored; extending just proximal to the terminus of the abdomen.
Abdomen: Widened, the fifth tergite the widest region before a gradual posterior narrowing; elliptical in shape. Tergites with expanded and triangular posterolateral tergal projections on the fifth through seventh segments. Supra-anal plate slightly transverse, a broadly rounded, blunt terminus, a small emargination present.
Head ( Fig. 41G): Transverse, the juxta-ocular protuberances small, but pronounced, the apex just lateral to the midline; the vertex is slightly convex, even with the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc, the entire carina depressed into a trough. 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 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; a central dark band running across the middle of the clypeus, the dorsal and ventral regions 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. 47N): A little less than three times as long as wide with a moderately defined supra-coxal bulge; dorsal surface entirely smooth or at most with very few, small tubercles. 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 numerous small tubercles; posterior margin with a medial emargination; the dorsal surface of the posterior third of the metazone not depressed. Mostly dark with pale and black markings across the surface, faint swirls present on the metazone just posterior to the supra-coxal sulcus.
Prothoracic Legs: Femur normal with a straight dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a very thin black band running medially from the base to terminus, some small segments more faded, two slight dark marks present dorsal to the band, one medial and the other near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the first two proximal posteroventral spines, in line with the most distal discoidal spine; pit is pigmented brown. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, 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 much 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 shorter or as long as the remaining segments combined.
Wings: Forewings mottled with brown, pale and black coloration; the costal region without defined banding distally, proximal region mostly brown with a green/pale anterior margin; vein coloration is pale, not matching surrounding cell coloration; a larger, but slightly more pale area is positioned centrally; brown color dominant across discoidal region with small regularly dispersed irregularly shaped black marks in the distal half. Forewings 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 the terminus of the abdomen. Hindwings with opaque discoidal region, colored rust or pale brown proximally and along the anterior margin, otherwise black; the anal region smoky black and translucent; the terminus of the discoidal region projecting far beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Slightly widened, the fifth or sixth tergite the widest region before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites with small posterolateral tergal projections on the terminal three segments. Supra-anal plate slightly wider than long, a broad, blunt terminus with a slight medial emargination. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51G.1-G.4): The main body of ventral left sclerite (L4A) broadly elliptical with rounded terminus, but with a rounded distal process (pda) positioned medially (may be short and rather blunt or more elongate and narrow), projecting at an angle towards the right phallomere (R1), appearing like a small, wellsclerotized 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) very short, quickly narrowing to a sharp point, not curved; the apical process (paa) elongate and thin, the terminus forming an angled blunt tip. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus; the ventral plate (pia) short with strongly defined grooves; the ventral process (pva) small and tooth-like and curved at the proximal base, the distal tip irregular, but narrowing to a point.
Head (Fig. 41H): Slightly transverse, the juxta-ocular protuberances large, the apex in the middle; the vertex is straight, higher than the dorsal margin of the eyes. The vertex and juxta-ocular protuberances evenly mottled with black, brown and pale markings.
Pronotum (Fig. 47O): As described for males. Prothoracic Legs: Femur normal with a near straight or slightly concave dorsal margin. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and just distal to the first most proximal posteroventral spine, in line with the most distal discoidal spine.
Meso-and Metathoracic Legs: As described for males. Wings: The costal region without defined banding, mostly pale with some brown mottling. The forewing extending just shy of the terminus of the abdomen.
Abdomen: Slightly widened, the fifth tergite the widest region before a gradual posterior narrowing. Tergites with small posterolateral tergal projections on the terminal four segments. Supra-anal plate almost square, a broad, blunt terminus with a slight medial emargination.
Etymology. A noun in the genitive case, L. kirtlandi is named in honor of Jared Potter Kirtland (1793-1877) for his contributions to natural science and medicine in northern Ohio as one of the founding trustees of the Cleveland Academy of Natural Science in 1845, later renamed the Kirtland Society of Natural History in his honor in 1865. The KSNH remained in existence until 1920 when the Cleveland Museum of Natural History was founded, the various natural history collections moving to the new museum. Diagnosis. One of the smaller species, L. manausensis is mostly black and whitish across the body with strongly asymmetrically colored forewings. Known only from north of Manaus, Brazil. The male genitalia are distinct from others in the Maya Group with its elongate ventral left sclerite (L4A) with a medial bulge.
Head (Fig. 42E): Transverse, the juxta-ocular protuberances small, but pronounced, the apex in the lateral half; the vertex is straight, just below the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc, the entire carina depressed into a trough, the margins sharp transitions. Ocelli small and protruding slightly on small cuticular mounds; the lateral ocelli oriented outward. The carina on the frons not pronounced, but present. Clypeus transverse, the upper margin slightly convex, the lower margin straight; the central, transverse carina not very pronounced, straight. Antennae scape and pedicel pale, the flagellum pale proximally, fading gradually to brown, then to black distally. Black band extending straight over the medial carina of the frontal suture, the center of the carina pale; two pale marks positioned on the lower region of the vertex. Frons pale; clypeus pale; the mandibles and labrum pale; the vertex and juxta-ocular protuberances mostly pale with brown speckling; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47P): A little less than three times as long as wide with a moderately defined supra-coxal bulge; dorsal surface with disperse, small tubercles in the posterior half. 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 with medial bulges; margins with numerous small tubercles; posterior margin with flat posterolateral corners and with a medial emargination; the dorsal surface of the posterior third of the metazone depressed. Mostly pale with black markings across the surface, especially just before the posterior margin and laterally just posterior to the supra-coxal sulcus.
Prothoracic Legs: Femur normal with a concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a three black bands, one basally, one medially that connects to the dorsal margin, and one adjacent to the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the first two proximal posteroventral spines, in line with the most distal discoidal spine; pit is pigmented brown. Posterior prothoracic femoral genicular spine half the length of the posteroventral spines, 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 much longer. Prothoracic coxae smooth, the anterior surface with 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 shorter or as long as the remaining segments combined.
Wings: Forewings mottled with pale (whitish) and black coloration; the costal region mostly pale with a few black marks, no regular banding; vein coloration is pale, not matching surrounding cell coloration; discoidal region evenly mottled with pale and black markings, whitish color more dominant distally. Forewings asymmetrically colored; one being mottled as described the other is darkened significantly with a black tone, the mottled pattern barely visible; extending just beyond the terminus of the abdomen. Hindwings with opaque black discoidal region; the anal region smoky black and translucent; the terminus of the discoidal region projecting far beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Slightly widened, the fifth or sixth 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 transverse with a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51H.1): The main body of ventral left sclerite (L4A) elongate, a rounded terminus with a medial bulge that could be considered as a slightly protruding and blunt distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate and forming a tapered sharp point with concave lateral margins, sometimes curved; the apical process (paa) elongate and thick, the terminus tapering to a narrowed, rounded tip. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus; the ventral plate (pia) long and broad with very well defined, curved grooves proximally; the ventral process (pva) small and tooth-like and curved at the proximal base, the distal tip irregular, but narrowing to a point.
Etymology. Named for Manaus, Brazil, near where the species was collected. 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 Taxonomic history. One of the earliest species to be described, L. maya was referenced in early works as M. annulipes, but this is likely due to the lack of comparison with the type of M. annulipes. Saussure and Zehntner recognized the species as unique, but as a variant of L. cayennensis. The species was later formalized as being distinct. It is apparent from the references that include L. 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 L. maya cannot be accurately confirmed or falsified and using historical records for biogeographic studies or population occurrence studies should be avoided.

Liturgusa maya
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 L. charpentieri was conducted in collections in Italy, but was never located (see Taxonomic History of L. guyanensis). However, the original description given by Giglio-Tos (1927) for L. charpentieri matches L. maya and with the expanded range of L. maya into South America as evidenced by specimens linking Central and South America through occurrences in Colombia, Venezuela, Ecuador and Peru, L. charpentieri can no longer be considered as distinct based on its southern distribution. Therefore, L. charpentieri Giglio-Tos, 1927, is now considered as a junior synonym to L. maya Saussure & Zehntner, 1894. Interestingly, L. 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 L. 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, L. 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 L. 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 L. 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, L. 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 L. 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 L. nubeculosa, which is a much larger species).
Diagnosis. Most similar to L. kirtlandi and L. trinidadensis with a similar size, coloration and pronotum shape, L. maya is distinct from the other two by a number of features including male genitalia. The most obvious difference easily diagnosing L. maya from L. 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, L. maya can be differentiated from L. trinidadensis by the larger apofisis falloid (afa) compared to the barely present, but sharp structure seen in L. trinidadensis. The species has the greatest size variation across its range. The main difference between L. maya and L. kirtlandi is the presence of tubercles on the pronotum in L. kirtlandi.
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.
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.

Diagnosis.
One of the largest species of Liturgusa, comparable to L. nubeculosa, L. stiewei is more comparable to many large species of Hagiomantis. Can be distinguished from L. nubeculosa by its shorter, more broad pronotum and black, more opaque hindwings.
Pronotum (Fig. 48A): A little less than three times as long as wide with a moderately defined supra-coxal bulge; dorsal surface smooth, without tubercles. Prozone broader than long, with parallel margins prior to rounded anterolateral corners, the anterior margin broadly straight; margins smooth or with very few broad, blunt tubercles. Metazone with concave lateral margins with near parallel margins just posterior to the middle; margins with numerous small tubercles; posterior margin with a shallow medial emargination; the dorsal surface of the posterior third of the metazone depressed. Coloration largely unknown except for two symmetrical dark marks on the anterior corners of the metazone.
Prothoracic Legs: Femur robust with a slightly concave dorsal margin; pale to dark banding not well defined on posterior (external) surface; anterior (internal) surface mostly not visible, but the distal half with a very thin black band running medially; the ventral surface pale. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the first two proximal posteroventral spines, in line with the most distal discoidal spine; coloration of pit not visible. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating at 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 longer. Prothoracic coxae not visible.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina present. Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings mottled with brown and pale coloration; the costal region with faintly defined banding; vein coloration is pale, not matching surrounding cell coloration. Hindwings are hidden in examined specimen.
Abdomen: Slender, the fifth tergite the widest region before a gradual posterior narrowing. Subgenital plate irregularly rounded and without styli.
Head (Fig. 42F): Longer than broad, the juxta-ocular protuberances very large, the apex in the middle; the vertex slightly concave or straight, above the dorsal margin of the eyes. Frontal suture without a carina, a black pigmented depression just ventral to medial region of suture, two depressed pits dorsally and symmetrically position lateral to the medial line that extend to broadly defined depressions dorsally towards the vertex. Ocelli small, all three of similar size and protruding on small cuticular mounds; the lateral ocelli oriented outward. The carina on the frons strongly pronounced, centrally elevated, the medial region just ventral to the carina strongly sloped ventrally. Clypeus transverse, the upper margin slightly straight medially and curving near dorsolateral corners; the lower margin concave or straight; the central, transverse carina pronounced and straight. Antennae scape pale, pedicel mostly pale with faint brown marks, the flagellum fading to black in the proximal quarter. Black markings extend laterally to the margin of the eyes from ventral depression abutting frontal suture. Vertex and juxta-ocular protuberances mostly pale with brown and black speckling; region between ocelli pale except for a thin band running along medial line from the frontal suture; lower region of frons pigmented brown; the dorsolateral corners of clypeus with brown spots, the rest is pale; the mandibles and labrum with pale and brown markings; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 48B): About 2.7 times long as wide with a moderately defined supra-coxal bulge; dorsal surface smooth, without tubercles. Prozone is broader than long with gradually tapering margins leading to an evenly rounded anterior margin; margins smooth. Metazone with concave lateral margins, the medial region flat or bulging outward; margins with very small tubercles; posterior margin with a very shallow medial emargination; the dorsal surface of the posterior third of the metazone depressed. Mostly pale or light brown with brown and black marking across the surface; two prominent black marks present in the posterolateral corners; two symmetrically positioned black marks on either side of midline and just anterior to the posterior margin; two symmetrically positioned lateral marks just posterior of supra-coxal sulcus.
Prothoracic Legs: Femur normal with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a strongly defined black band running medially from the base to terminus, some regions slightly widened, particularly near the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A large femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the first two proximal posteroventral spines, in line with the most distal discoidal spine; pit is black. Posterior prothoracic femoral genicular spine tiny compared to posteroventral spines, originating at the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) and fourth through sixth of similar length, the second and third longer. Prothoracic coxae smooth, the anterior surface with a large medially positioned black band in the proximal half as well as a similar black band in the distal half.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina present. Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings mottled with brown, pale and black coloration; the costal region mostly pale with some regularly spaced black marks; vein coloration pale; discoidal area with highly contrasting regions, evenly mottled proximally with a distinct shift to mostly pale in the distal half. Forewings symmetrically colored; extending just short of the terminus of the abdomen, the supra-anal plate visible. Hindwings entirely opaque black; the terminus of the discoidal region projecting beyond the distal margin of anal region, the wing appearing slightly elongate.
Abdomen: Widened, the fourth tergite the widest before a gradual posterior narrowing; a smooth surface with light brown coloration. Tergites without posterolateral tergal projections, if present they are very small. Supra-anal plate transverse, an evenly rounded terminus. Diagnosis. Very similar to L. maya in size and shape, but with an even color patterning on the forewings, lacking highly contrasting regions. The hindwings for L. tessae are very rusty colored in males and pale to rusty colored in females along the anterior half of the discoidal region. The male genitalia are distinct in that the ventral left sclerite (L4A) is elongate, the terminus tapers narrowly before giving rise to a short, blunt distal process (pda).
Frontal suture with a slight medial carina forming a continuous arc. Ocelli small, the central slightly larger than lateral two, all three protruding on small cuticular mounds; the lateral ocelli oriented outward. The carina on the frons thin, 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 fading to black in the proximal quarter. Black band extending straight over the medial carina of the frontal suture, 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 medial region of the vertex. Lower region of frons pigmented brown; the dorsolateral corners of clypeus with brown spots, the rest is pale; the mandibles and labrum with pale and brown markings; the vertex and juxta-ocular protuberances mostly pale with brown or black speckling; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 47S): About 2.5 times long as wide with a moderately defined supra-coxal bulge; dorsal surface with very small, disperse tubercles in the posterior half. Prozone a little broader than long with near parallel margins that gradually taper to an evenly rounded anterior margin; margins with few blunt tubercles. Metazone with concave lateral margins, the medial region flat or slightly bulging outward; margins with small tubercles; posterior margin with a shallow medial emargination; the dorsal surface of the posterior third of the metazone depressed. Mostly pale or light brown with brown and black markings across the surface, two prominent black marks present in the posterolateral corners.
Prothoracic Legs: Femur normal with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a black mark near the base and a medial black band beginning in the distal half and terminating near the end of the femur; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and between the first two proximal posteroventral spines, in line with the most distal discoidal spine; pit is pale. 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 black mark medially in the proximal half.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina present. Mesotarsi with first segment as long or slightly shorter than the remaining segments combined.
Wings: Forewings mottled with brown, pale and greenish coloration; the costal region without strongly defined banding, mostly pale with dispersed brown markings; vein coloration matching the corresponding surrounding cell colors; lacking noticeable central or proximal pale markings, the wing having evenly mottled coloration that is not highly contrasting. Forewings asymmetrically colored; one being mottled as described the other is darkened with a rust tone, the mottled pattern still visible; extending just beyond the abdomen. Hindwings rust colored (more pale proximally), the discoidal region more opaque, but translucent near the boundary with the anal region; the anal region translucent and rusty colored; 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, light brown or rust coloration. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 51J.1-J.6): The main body of ventral left sclerite (L4A) elongate, the terminus tapering to a narrow point; a short, blunt distal process (pda) is centrally positioned; the right margin in the distal quarter highly sclerotized, extending to the terminus of the pda; an elliptical depression on left half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) short, quickly narrowing to a sharp point, the lower margin concave and often irregular, leading to a secondary process that is short and rounded; the apical process (paa) broad and elongate, tapering to a narrow and rounded terminus. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) broad, tapering slightly to a rounded, blunt and membranous terminus; the ventral plate (pia) long, broad proximally with strongly defined grooves; the ventral process (pva) enlarged, c-shaped with a smooth surface, the distal tip broad and rounded.
Head (Fig. 42D): Slightly transverse, the juxta-ocular protuberances large, the apex just lateral of the middle; the vertex concave, above the dorsal margin of the eyes. Ocelli positioned on a small carina connecting all three. Antennae scape and pedicel pale, the flagellum fading to black by the middle. In addition to the black band extending straight over the medial carina, the anterior face, vertex and juxta-ocular protuberances mottled with pale, brown and black markings. Lower region of frons pigmented brown or light brown; the clypeus, mandibles, and labrum with pale and brown markings.
Pronotum (Fig. 47T): As described for males. Prothoracic Legs: Anterior (internal) surface of femur with a black mark near the base and a medial black band that may be interrupted or degraded beginning in the distal half and terminating near the end of the femur. A shallow femoral pit that is pigmented brown or pale.
Meso-and Metathoracic Legs: As described for males. Wings: Forewings may be asymmetrically colored like in males; extending almost to the terminus of the abdomen, but the supra-anal plate still visible. Hindwings smoky colored with a rusty base and anterior margin, opaque; the terminus of the discoidal region projecting a little beyond the distal margin of anal region, the wing appearing moderately elongate.
Abdomen Diagnosis. Most similar to L. maya and L. kirtlandi, exhibiting similar size, coloration and pronotum shape, L. trinidadensis is distinct from the other two by a few characteristics. First, the supra-anal plate in both males and females is nearly square with a broad and blunt terminus. Second, the central ocellus is the same size as the lateral two while L. maya and L. kirtlandi have larger central ocelli. Finally, L. trinidadensis has numerous tubercles in the posterolateral corners of the metazone. The species is also entirely restricted to the island of Trinidad, not extending into mainland South America.
Head (Fig. 43A): 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 ocellus the same size as the lateral, all are 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 sloped. Clypeus transverse, the upper margin convex, the lower margin concave; the central, transverse carina pronounced and straight. Antennae mostly pale, fading to a darker brown distally. Black band extending straight over the medial carina of the frontal suture, pale medially; two black bands lateral to the lateral ocelli; two black marks just dorsal to frontal suture. Black band extending across the lower region of the frons, but separate from the ventral margin; brown markings in the dorsolateral corners for the clypeus that extend medially along the central carina; the mandibles and labrum with pale and brown markings; the vertex and juxta-ocular protuberances mostly pale with fine, disperse black speckling; the area immediately adjacent to ocelli black. Palpi are pale.
Pronotum (Fig. 48C): A little less than three times long as wide with a moderately defined supra-coxal bulge; some tubercles in the posterior third, but otherwise 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, the nadir of the margins in the posterior half almost parallel for a short distance before widening to the posterior margin; margins with small tubercles; posterior margin with a medial emargination; the dorsal surface of the posterior third of the metazone slightly depressed; tubercles present in the posterolateral corner. 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 without a black band, but three thin dashes positioned medially, one in the proximal third, one in the middle and one next to the femoral brush; the ventral surface pale. Posterior surface of femur with few tubercles. A shallow femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and slightly distal to the first most proximal posteroventral spine, in line with the most distal discoidal spine; pit is pigmented pale or brown. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third longer. Prothoracic coxae smooth, the anterior surface with a very small, black mark medially in the proximal half as well as a small, circular 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 without defined banding proximally, mostly brown and green mottling; the distal half of the costal region with regular banding; 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 larger pale area is positioned centrally. Forewings often, but not always asymmetrically colored; one being mottled as described the other is slightly darkened, the mottled pattern still visible; extending just beyond 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 square with rounded posterolateral corners and a blunt terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52A.1-A.5): The main body of ventral left sclerite (L4A) with rounded terminus, but with a distal process (pda) positioned just lateral to the middle that is short and tapering to a sharp point, projecting at an angle laterally, appearing like a small, well-sclerotized tooth (tooth absent in one examined specimen); 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, barely present, quickly narrowing to a tiny, sharp point; the apical process (paa) elongate and thin, the terminus evenly rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus, the end often folded; 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.
Head (Fig. 43B): About as long as broad, the juxta-ocular protuberances large, the apex just lateral to the midline; the vertex is straight, but with two bulges just medial to the parietal sutures, slightly higher than dorsal margin of the eyes. Ocelli raised slightly on a continuous carina connecting all three. The carina on the frons pronounced, the medial region just ventral to the carina sloped. Clypeus transverse, the upper margin convex, the lower margin concave; the central, transverse carina pronounced and straight, the ventral half depressed. The vertex and juxta-ocular protuberances mostly dark brown with fine pale speckling.
Pronotum (Fig. 48D): Tubercles in the posterior half, but otherwise smooth; numerous tubercles present in the posterolateral corner of metazone. Two lateral bulges in the dorsal surface around the midline of the metazone.
Prothoracic Legs: Prothoracic tibial posteroventral spines with the first (proximal) smallest and the third through sixth of similar length, the second longer.
Meso-and Metathoracic Legs: As described for males. Wings: The costal region of forewing without defined banding, mostly brown and green mottling. Forewings are darker brown overall than in males; far shorter than the terminus of the abdomen, often terminating around the sixth segment. Hindwings with the terminus of the discoidal region projecting slightly beyond the distal margin of anal region.
Abdomen: Slightly widened, the fifth tergite the widest region before a gradual posterior narrowing. Seventh tergite with small posterolateral projections. Supra-anal plate almost square, a broad, blunt terminus with a slight medial emargination.
Etymology. Named for the island of Trinidad, where this species inhabits and appears to be endemic. Diagnosis. A medium sized species, is distinct from all other Liturgusa. Could be considered most similar to L. guyanensis based on the pronounced pronotal shape modifications compared to all other Liturgusa, but L. zoae has a longer pronotum. The prothoracic femora lacks tubercles on the posterior (external) surface, unique to the species. The hindwings are dark black and highly opaque, appearing rounded.
Head (Fig. 43C): Transverse, the juxta-ocular protuberances prominent, the apex in the lateral half; the vertex is concave, dipping just prior to the parietal sutures, even with the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, the region just ventral and dorsal to the carina depressed, the carina within a trough. Ocelli small, all the same size, protruding and laterally angled on a pronounced, curved carina that connects all three. The carina on the frons pronounced, the medial region just ventral to the carina depressed. Clypeus transverse, the upper margin convex, the lower margin slightly convex, the lateral margins adjoining lower margins with rounded corners; the central, transverse carina pronounced and slightly curved, two small grooves symmetrically located just dorsal to the carina. Antennae scape pale, pedicel dark brown, the flagellum fading to dark brown just slightly distal to the base. Vertex and juxta-ocular protuberances mottled pale and brown; cuticle just adjacent to ocelli dark brown or black; pale centrally to the three ocelli. Frons, clypeus, labrum, mandibles and palpi pale. Pronotum (Fig. 48E): A little less than three times long as wide with a defined supra-coxal bulge; dorsal surface without tubercles; the lateral margins expanded and slightly lamellar. Prozone slightly broader than long with pronounced convex lateral margins; the anterior margin round; the dorsal surface spherical or bulbous, raised higher than seen in other Liturgusa; lateral margins smooth or with very few blunt tubercles. Metazone with concave lateral margins, tapering rapidly posterior of the supra-coxal sulcus, the medial region bulging slightly outward before widening to a rounded posterior margin; posterior margin with a small emargination; lateral margins with disperse tubercles, mostly located in the anterior third; the dorsal surface of the posterior third of the metazone very depressed; symmetrical posterior bulges pronounced. Mostly pale with black markings, two prominent black marks near the posterior margin of the metazone and laterally just posterior of 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 thinned or interrupted medially, the band thicker near the femoral brush and connected to the dorsal margin; the ventral surface pale. Posterior surface of femur without tubercles. A femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and in line with the first most proximal posteroventral spine, proximal to the most distal discoidal spine; pit is pigmented black. Posterior prothoracic femoral genicular spine smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Ventral surface with raised, blunt carina just medial to the posteroventral spines beginning at the femoral pit and fading to flat after the third most distal posteroventral spine. Prothoracic tibial posteroventral spines with the first (proximal) smallest, the fourth and fifth slightly longer and the second, third and sixth of similar length. Prothoracic coxae smooth, the anterior surface with a large black band medially in the proximal half as well as a 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 shorter or at most as long as the remaining segments combined.
Wings: Forewings mottled with brown, whitish pale, and black coloration; the costal region mottled matching the discoidal region, some banding pattern present medially; the costal region widened; vein coloration across discoidal region pale or light brown; a large pale area is positioned centrally; most of the surface dark brown or black with whitish mottling across the surface, the distal tip more whitish. Forewings asymmetrically colored; one being mottled as described the other is darkened significantly with a rust tone, the mottled pattern still visible; extending just beyond the abdomen. Hindwings opaque black, the veins black; the terminus of the discoidal region projecting just beyond the distal margin of anal region, the wing appearing rounded.
Abdomen: Slightly widened in the middle, the fourth or fifth tergite the widest before a gradual posterior narrowing; a smooth, brown and black colored dorsal surface. Tergites with tiny triangular posterolateral tergal projections. Supra-anal plate transverse, an evenly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52B.1): The main body of ventral left sclerite (L4A) elongate, an evenly rounded terminus with the margin rolled slightly along the terminus; the left side with an elongate depression on the surface; lacking a distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) forming a very large triangular projection that evenly tapers to a point; the apical process (paa) elongate and thin, the terminus rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers rapidly to a narrowed and rounded, membranous terminus; the ventral plate (pia) long, curved with a medial tooth projecting towards the pva, smooth surface; the ventral process (pva) long and smooth, but with a sharply bent terminus, creating a notch.
Head (Fig. 43D): About as long as broad, the juxta-ocular protuberances prominent, the apex in the middle; the vertex is slightly concave, evenly sloping to a slightly depressed center, above the dorsal margin of the eyes. Region just ventral to the frontal suture slightly depressed. Ocelli very small, all the same size, the lateral ocelli widely positioned, almost in line with the middle of the antennal insertion. The carina on the frons present, but not strongly pronounced. Antennal scape pale, pedicel with black markings, the flagellum brown proximally, fading darker and darker distally. Cuticle just adjacent to ocelli pale or light brown. Pronotum (Fig. 48F): Less than three times long as wide with a defined supra-coxal bulge. Posterior margin of metazone with flat posterolateral corners and with a small medial emargination; lateral margins with numerous, pronounced tubercles.
Prothoracic Legs: Prothoracic tibial posteroventral spines with the first (proximal), third through fifth short, the second and sixth longer and similar length..
Meso-and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings with the costal region mottled matching the discoidal region, some banding pattern distally; the costal region widened and extending distally; vein coloration mostly matches surrounding coloration; 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; black with whitish pale spots in the distal half.
Forewings asymmetrically colored; one being darker and less contrasting; extending short of the terminus of the abdomen.
Abdomen: Slightly widened, the fourth or fifth tergite the widest region before a gradual posterior narrowing. Tergites with posterolateral projections of varying size, but shaped like expanded triangles and not posteriorly oriented teeth. Supra-anal plate transverse, an evenly rounded terminus.
Etymology. A noun in the genitive case, L. zoae is named for my daughter Zoey Kay Svenson.
Interestingly, the species was included in a number of studies focused on chromosomes headed by Sally Hughes-Schrader in the 1940's and into the 60's. Other than being included in species records for regional studies and taxonomic lists, the species has received no taxonomic attention since its original description.
Natural history. Encountered commonly in lowland tropical forest in southern Nicaragua. Individuals were present on trees in agricultural fields, secondary forest as well as primary forest regions, but with lower densities. Usually found positioned head down on the tree trunk and varying elevations. Individuals ran rapidly up and/or to the opposite side of the trunk when approached. All trees were smooth bark. Found living sympatrically with L. fossetti, L. maya, and C. atricoxata.
Diagnosis. A small, slender species like L. dominica and L. milleri, but restricted in distribution to Central America. The forewings of males are colored asymmetrically while L. dominica and L. milleri both have symmetrically colored forewings; the costal region in males and females with a contrasting green color in newly preserved specimens, but usually faded to pale in older specimens. The genitalia of males are unique among Liturgusa with an elongate and narrowed distal terminus of the right dorsal phallomere (fda) of the first sclerite of right phallomere (R1).
A thorough description of the species was provided by Rehn (1950) and can be referred to in addition to the standardized description provided herein.
Head (Fig. 43E): Slightly transverse, the juxta-ocular protuberances small, the apex in the lateral half; the vertex between the parietal sutures with two convex regions, the medial line being slightly depressed; vertex slightly lower than the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and positioned on the edge of a blunt, curved carina that connects all three ocelli, continuing slightly beyond the lateral ocelli. Frons transverse, the lower portion narrowed under the antennal insertions. Lateral ocelli oriented outward. Upper margin of clypeus convex, lower margin straight; a medial transverse ridge moderately pronounced, rounded. Antennae pale basally, fading almost immediately to black. Two black spots lateral to the frontal suture at the base of each parietal suture; lower region of frons with a broad black band; the clypeus mostly pale with two brown spots in the upper corners; the mandibles pale with lateral darkening; the labrum with a pale upper region and a brown to black lower region; the vertex and juxta-ocular protuberances with brown and black markings; thin, black margins surround the ocelli. Palpi are pale.
Pronotum (Fig. 48G): Elongate with a defined supra-coxal bulge; dorsal surface smooth. Prozone elongate with gradually widening margins before tapering anteriorly to a rounded anterior margin; the lateral margins smooth. Metazone with concave lateral margins, narrowing quickly posterior to the supra-coxal bulge to the midpoint, then widening gradually until reaching the rounded posterior terminus; margins with few small tubercles; posterior margin barely medially emarginate; the dorsal surface of the posterior third of the metazone barely depressed. Pale coloration dominant with strong black marks near the anterior and posterior margins and symmetrical black swirls on the supra-coxal bulge.
Prothoracic Legs: Femur elongate 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, often interrupted medially; a broad black mark at the base and just distal to the tibial spur groove; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medial to the most proximal posteroventral spine; pit is pale. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are longer (the second being the longest). Prothoracic coxae smooth; the anterior surface with a small, black band medially in the proximal half as well as a black spot medially towards the distal terminus.
Meso-and Metathoracic Legs: Femora with a faint ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment as long or longer than remaining segments combined.
Wings: Forewings mottled with pale, brown, and green coloration; the costal region with pale/green and dark near regular banding; the discoidal region evenly mottled with large brown and pale color markings. Forewings often colored asymmetrically, one being mottled as described above while the other is rust colored with the mottled pattern still visible; extending to around the terminus of the abdomen. Hindwings smoky black, but translucent, the discoidal region darker and more opaque; the discoidal region narrowed; the costal region dark brown proximally; the terminus of the discoidal region projecting slightly beyond the distal margin of anal region.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; a smooth brown and black colored dorsal surface. Supra-anal plate highly transverse, tapering quickly to a broad, blunt and wide terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52C.1-C.3): The main body of ventral left sclerite (L4A) with a narrowed, but rounded terminus, lacking a distal process (pda); an elliptical depression on the lateral half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate and finger-like, margins narrowing before reaching an expanded and bulbous terminal end, either straight or curved; the apical process (paa) short, curved and with a rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers dramatically to a narrowed and membranous terminus, almost pointed, but the tip is blunt; the ventral plate (pia) short and toothlike; the ventral process (pva) c-shaped and smooth.
Head (Fig. 43F): As broad is high, the juxta-ocular protuberances large, the apex in the lateral half; the vertex between the parietal sutures concave; vertex slightly higher than the dorsal margin of the eyes. Frontal suture with a medial carina forming a high and continuous arc. Lateral ocelli oriented anterolaterally. Clypeus slightly transverse, upper margin convex, lower margin straight. Antennae pale basally, fading almost immediately to black. Black markings surround frontal suture, are present on the vertex and juxta-ocular protuberances. Lower region of frons with a broad black band; the mandibles pale with distal darkening; the labrum mostly pale with a brown medial strip.
Pronotum (Fig. 48H): Margins of prozone with few small tubercles. Prothoracic Legs: As described for males. Meso-and Metathoracic Legs: As described for males. Wings: Forewings colored symmetrically; shorter than the abdomen, reaching the fifth or sixth tergite. Hindwings with the costal region light brown proximally; this distal margin of the discoidal region pale or light brown.
Abdomen: Widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supra-anal plate highly transverse, evenly rounded. Diagnosis. This small species can be identified based on its slender size and its geographic restriction to the Caribbean island of Dominica. Much like L. milleri, the forewings also have white or grey spots in proximal quarter just posterior to the radial vein. The forewings are also net-like in appearance with pale veins and black pigmented cells. The anterior surface of the prothoracic coxae has no pigmentation, while L. milleri and L. cursor both have black markings. In addition, the juxta-ocular protuberances are very small and the entire vertex is concave, falling well below the dorsal margin of the eyes in both sexes. The other two species that resemble L. dominica are restricted to French Guiana and Central America (L. milleri and L. cursor, respectively).
Head (Fig. 43G): Transverse, the juxta-ocular protuberances barely present, reduced to a slight bulge, the apex of which is in the lateral half; the vertex is concave overall; the medial region well below the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, but depressed into the anterior surface of the head, more so ventral to the suture. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward, a few degrees off perpendicular. Clypeus transverse, the upper margin convex, the lower margin straight. Antennae pale at the base, the flagellum fading to brown near the base. Thin black band extending straight over the medial carina of the frontal suture, the medial portion of the carina pale; lower region of frons with a black band; the clypeus is pale; the mandibles and labrum mostly pale, but with some brown marks; the vertex and juxta-ocular protuberances pale, but with four brown and black marks positioned on each juxta-ocular protuberance and between the parietal sutures; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 48I): Highly elongate with a defined supra-coxal bulge; dorsal surface entirely smooth. Prozone elongate with slightly convex lateral margins that taper anteriorly; the margins smooth. Metazone with strongly concave lateral margins, a slight bulge in the posterior half; margins with small tubercles; posterior margin with a medial emargination; the dorsal surface of the posterior third of the metazone barely depressed. Pale with strong black marks across the surface, swirls present at the supracoxal bulge.
Prothoracic Legs: Femur elongate with a slightly concave dorsal margin; tubular in overall shape with the dorsal margin less defined, the anterior and posterior surfaces almost continuous; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a black band running medially from the base to terminus; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medial to and exactly between the first two proximal posteroventral spines, but slightly proximal to the most distal discoidal spine; pit is pigmented darkly. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating just distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are much longer (about half the length of the terminal spine). Prothoracic coxae smooth, the anterior surface pale.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment as long or shorter than remaining segments combined.
Wings: Forewings mottled with black, light brown, and greenish coloration; the costal region without strong banding, pale and black proximally and mostly black distally; veins are pale and cells are black across the discoidal and distal portion of the costal region, giving a contrasting net-like appearance; two pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein. Forewings colored symmetrically; extending just beyond the terminus of the abdomen. Hindwings opaque brown, the discoidal region more pale proximally; the costal region light brown proximally and narrowing to the anterior margin distally; the terminus of the discoidal region barely projecting beyond the distal margin of anal region, the distal margin strongly emarginate between the anal and discoidal region.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; a smooth, brown and black colored dorsal surface. Supra-anal plate transverse, tapering gradually to a rounded terminus; the terminus with a slight emargination. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52D.1): The main body of ventral left sclerite (L4A) with rounded terminus, but often with a slight bulge just lateral to the medial line, lacking a distal process (pda); sometimes a depression on the lateral half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate and robust, tapering to a strong point and angled off the central axis of the L4B; the apical process (paa) broad, shortened, cylindrical and curved, the terminus a 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 and irregular, with grooves; the ventral process (pva) smooth and tapering to a point distally, one edge straight and the other convex, tooth-like in appearance.
Head (Fig. 43H): Head is transverse, the juxta-ocular protuberances very small, reduced to a bulge, the apex of which is in the lateral half; the vertex is concave overall, the medial region below the dorsal margin of the eyes. Clypeus transverse, the upper margin convex, the lower margin concave. Antennae pale at the insertion, fading to a brown black at the first antennomere. The clypeus is pale with two dorsolateral dark spots; the mandibles and labrum mostly pale, but with some brown marks; the vertex and juxta-ocular protuberances pale, but with disperse brown markings; the area immediately adjacent to lateral ocelli black. Palpi are pale.
Meso-and Metathoracic Legs: As described for males. Wings: Forewings shorter than the abdomen, often terminating at the fifth tergite. Hindwings with the terminus of the discoidal region barely projecting beyond the distal margin of anal region, the distal margin strongly emarginate between the anal and discoidal region.
Abdomen: Widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, rounded.
Ootheca (Fig. 21A-B). Like all other known oothecae of Liturgusa, L. dominica produces a broad based case with a narrowing tube extending dorsolaterally away from the substrate. The eggs are contained within the bulbous base that is almost spherical excluding the flattened margin where the case is attached to the bark of a tree or branch. The tube narrows quickly and is short, about half the diameter of the main body. The tube is hollow and the entry is flat, but at an angle to the long axis of the tube, the upper margin extending further than the lower margin forming an elliptical opening that is parallel to the substrate (may reduce water intrusion).
Etymology. A noun in apposition, L. dominica is named for the island of Dominica, where this species inhabits and appears to be endemic. Diagnosis. This small species can be identified based on its slender size, its geographic restriction to the Guianas as well as the two symmetrically located white spots in the proximal quarter of their forewings. The other two species that resemble L. milleri are restricted to the island of Dominica and Central America (L. dominica and L. cursor, respectively).
Head (Fig. 43I): Transverse, the juxta-ocular protuberances small, the apex in the lateral half; the vertex between the parietal sutures is slightly concave, uneven; vertex lower than the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, but depressed into the anterior surface of the head. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward, a few degrees off perpendicular. Clypeus highly transverse, the upper margin convex, the lateral and lower margin forming a continuous rounded margin. Antennae pale at the base, the scapes with two dark marks, the flagellum fading to black within a few antennomeres from the base. Broad black band extending straight over the medial carina of the frontal suture, the carina remaining white; two white spots between the lateral ocelli; lower region of frons with a black band; the clypeus with two black spots in the upper lateral corners; the mandibles and labrum pale; the vertex and juxta-ocular protuberances pale, but with four black marks, two positioned on each side of the parietal sutures; the area adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 48K): Highly elongate with a defined supra-coxal bulge; dorsal surface entirely smooth. Prozone elongate with slightly convex lateral margins that taper anteriorly; the margins smooth. Metazone with strongly concave lateral margins, a slight bulge in the posterior half; margins with numerous small tubercles; posterior margin with a strong medial emargination; the dorsal surface of the posterior third of the metazone barely depressed. Pale with strong black marks across the surface, swirls present at the supra-coxal bulge.
Prothoracic Legs: Femur elongate with a slightly concave dorsal margin; tubular in overall shape with the dorsal margin less defined, the anterior and posterior surfaces almost continuous; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a black band running medially from the base to terminus; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medial and just distal to the first most proximal posteroventral spine and in line with the distal most discoidal spine; pit is black. Posterior prothoracic femoral genicular spine smaller than all other posteroventral spines, originating just proximal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are longer. Prothoracic coxae smooth; the anterior surface with a broad, black band medially in the proximal half as well as a black spot medially towards the distal terminus.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment longer than remaining segments combined. Wings: Forewings mottled with pale, black, and brown coloration; the costal region with pale or whitish and dark irregular banding; two bright pale spots are positioned in the proximal quarter of the discoidal region just posterior to the first radial vein. Forewings colored symmetrically; extending just beyond the terminus of the abdomen. Hindwings smoky black, but translucent, the discoidal region darker and more opaque; the costal region pale proximally and narrowing to the anterior margin distally, the pale margin continuing across the terminal margin of the discoidal region; the margin of the anal region black; the terminus of the discoidal region barely projecting beyond the distal margin of anal region, the distal margin strongly emarginate between the anal and discoidal region.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; a smooth and black colored dorsal surface. Supra-anal plate transverse, tapering quickly to a blunt and wide terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52E.1): The main body of ventral left sclerite (L4A) with a slight medial bulge, but mostly a rounded terminus, lacking a distal process (pda); sometimes a depression on the lateral half is present. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) elongate and robust, tapering to a strong point, one margin being concave and the other being convex; the apical process (paa) broad, shortened, cylindrical and curved, the terminus with a rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus, both margins sclerotized with a broad central membranous gap; the ventral plate (pia) smooth and broad; the ventral process (pva) smooth and tapering to a point distally, one edge straight and the other convex.
Head (Fig. 43J): Juxta-ocular protuberances large, the apex in the middle; the vertex between the parietal sutures is concave; vertex even or just higher than the dorsal margin of the eyes. Lower region of frons with black marks laterally and a central black mark, two pale gaps on each side of the central black mark; the mandibles pale with brown markings distally.
Prothoracic Legs: Prothoracic coxae smooth; the anterior surface with a broad, black band medially in the proximal half as well as a black spot medially in the distal half.
Meso-and Metathoracic Legs: Mesotarsi with first segment as long or slightly longer than remaining segments combined.
Wings: Forewings shorter than abdomen, often terminating prior to the narrowing of the abdomen.
Abdomen: Widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supra-anal plate slightly transverse, rounded.
Etymology. A noun in the genitive case, L. milleri is named for Kelly B. Miller for his contributions to Mantodea sampling and his valuable collaboration.

Cursor Group B
Liturgusa actuosa Rehn, 1950 http://species-id.net/wiki/Liturgusa_actuosa Liturgousa actuosa : Rehn 1950: 377-382, Figs 12-17;Hughes-Schrader 1951: 178, 183-184, 186-187, Tables 1-2, Fig. 2;Hughes-Schrader 1953: 544-554;Callan and Jacobs 1957: 201;Otte 1978: 76;Liturgousa arcuosa: Hughes-Schrader 1950: 11-14, 38-39,  Taxonomic history. Described in 1950 by James Rehn, the species was only known from Barro Colorado Island in the Republic of Panama. At the time, it was considered most similar to L. annulipes rather than the sympatric L. cursor, but this study establishes that L. annulipes is not distributed in Central America and L. actuosa was being compared with an unknown taxon considered as L. annulipes at the time. Interestingly, the species was included in a number of studies focused on chromosomes headed by Sally Hughes-Schrader in the 1940's and into the 60's. Other than being included in species records for regional studies and taxonomic lists, the species has received no taxonomic attention since its original description.
Diagnosis. The species is most similar in appearance to L. cameroni and L. nubeculosa, but is much smaller and restricted entirely to Central America. The pronotum is moderately elongate and the coloration of forewing is more evenly brown rather than exhibiting the highly contrasting mottled color patterns as seen in L. nubeculosa. The costal region of the forewing is more pale or green with black markings that are not consistent with regular banding. In addition, the discoidal region of the hindwing projects well beyond the terminal margin of the anal region while the hindwing of L. cameroni and L. nubeculosa are more truncate. The wings extend just shy of the tip of the abdomen in females, but extend slightly beyond in males.
A thorough description of the male and female was provided by Rehn (1950) and can be referred to in addition to the standardized description provided herein.
Pronotum (Fig. 49A): Moderately elongate with a slightly defined supra-coxal bulge; dorsal surface with very few, very small tubercles that are mostly in the posterior half of the metazone. Prozone with lateral margins that are gently convex, tapering anteriorly; the margins smooth or at most with one or two very small tubercles. Metazone with concave lateral margins; margins with numerous small tubercles; posterior margin medially emarginate; the dorsal surface of the posterior third of the metazone not depressed. Brown coloration dominant, but a few strong black marks.
Prothoracic Legs: Femur elongate with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface, but dark areas are dominant; area between the posteroventral spines black; anterior (internal) surface with a thin black band running medially from the base to terminus; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is entirely black. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) and fourth through sixth approximately the same size, the second and third being much longer. Prothoracic coxae smooth; the anterior surface with a small black mark in the proximal half positioned medially in both orientations.
Meso-and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina obvious. Mesotarsi with first segment at most equal to remaining segments combined.
Wings: Forewings mostly brown with darker splotching, the anterior margin of the costal region green or pale, the veins are mostly pale; lacking highly contrasting coloration and more evenly dark brown; the costal region with dark marks, but not regularly banded. Forewings colored symmetrically. Hindwings smoky black with the anterior and basal region of the discoidal region with faded dark brown coloration; the anal region smoky and translucent; the terminus of the discoidal region projecting well beyond the distal margin of anal region, the wing elongate in appearance.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; smooth, a brown and black colored dorsal surface. Supra-anal plate transverse, tapering to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli. Genital Complex (Fig. 52F.1-F.3): The main body of ventral left sclerite (L4A) with a small, dull, cone-shaped distal process (pda) positioned laterally on the terminus and oriented 45 degrees from the central axis of the L4A. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) straight and tapering into a short, sharp point; the apical process (paa) cylindrical and curved, the terminus evenly rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus; the ventral plate (pia) strongly sclerotized and narrow, with a few strongly defined and curved grooves; the ventral process (pva) small and irregularly shaped, tapering to a point with a rough surface oriented towards the pia.
Head (Fig. 44B): Juxta-ocular protuberances moderately pronounced, the apex in the middle third; the vertex between the parietal sutures is slightly concave; vertex well above the dorsal margin of the eyes. Ocelli small and laying nearly flat. Antennae pale brown basally, fading gradually to dark brown. Clypeus mostly pale with lateral and lower margins that are dark brown, but pale on the very edge; the labrum mostly pale; the vertex and juxta-ocular protuberances brown with disperse black splotches. Palpi are pale.
Pronotum (Fig. 49B): Dorsal surface with few, very small tubercles. Prothoracic Legs: Femur moderately elongate with a slightly concave dorsal margin. Meso-and Metathoracic Legs: As described for males. Wings: As described for males. Abdomen: Widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supra-anal plate almost as long as wide, evenly rounded. Natural history. Males and females found in local abundance at the Madre Selva Biological Research Station in the Loreto Province, Peru. The species was living in sympatry with L. krattorum on the same smooth bark, medium diameter trees. The species was easily collected during the day and often found at reachable heights in lower sections of the tree.
Diagnosis. Extremely similar in coloration, size, and distribution to L. krattorum. Posterior prothoracic femoral genicular spine of female tiny, much smaller than seen in any other species. Overall color mottling darker with greater contrast, the hindwings being darkly smoke colored or nearly black and never rusty as in L. krattorum.
Head (Fig. 44C): Juxta-ocular protuberances large, the apex in the lateral third; the vertex between the parietal sutures is straight; vertex just below the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward, a few degrees off perpendicular. Upper margin of clypeus convex, lower margin slightly concave with a medial bulge. Antennae pale basally fading quickly to black. Broad black band extending from eye to eye over the medial carina of the frontal suture; lower region of frons with a transverse black band; vertex and juxta-ocular protuberances speckled with brown and black marks. Palpi are pale.
Pronotum (Fig. 49C): Elongate with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone with lateral margins that are parallel, tapering anteriorly; the margins smooth. Metazone with concave lateral margins; margins smooth or at most with very small and blunt tubercles; posterior margin medially emarginate; the dorsal surface of the posterior half slightly depressed, evenly concave.
Prothoracic Legs: Femur elongate 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; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is colored black. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating at the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the second through sixth of similar length, the second and third may be slightly longer. Prothoracic coxae smooth; the anterior surface with a medial black band centrally located in the central 75% of the surface; a very small black spot on the distal lobe.
Wings: Forewings mottled with brown, black, and pale coloration; the costal region with light to dark irregular banding. Forewings colored asymmetrically, one being mottled the other is blackened with the mottled pattern still slightly visible. Hindwings smoky black with rusty coloration at the very base of the discoidal region; the terminus of the discoidal region projecting well beyond the distal margin of anal region.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; smooth surface. Supra-anal plate transverse, tapering to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52G.1-G.2): The main body of ventral left sclerite (L4A) with a prominent, distal process (pda) positioned medially and tapering to a sclerotized point that is oriented in line with the central axis of L4A; the margins of the pda sclerotized and either straight or convex. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) tapering to an elongate, sharp point that is well sclerotized and oriented approximately 35 degrees from the central axis; the apical process (paa) cylindrical and gently curved, the terminus being an expanded, blunt knob. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus, the lateral margins being sclerotized and robust with a medial, membranous gap at the terminus; the ventral plate (pia) strongly sclerotized, broadened proximally with large curved grooves; the ventral process (pva) c-shaped, broad, and rounded distally, the surface slightly rough.
Head (Fig. 44D): The vertex between the parietal sutures is slightly concave; vertex just above the dorsal margin of the eyes. Ocelli small and laying flatly on the surface. Lower margin of clypeus straight with a slight medial bulge. Broad black band extending from eye to eye over the medial carina of the frontal suture, but with medial gap; lower region of frons with lateral black markings, a pale medial region. Palpi darkened terminally.
Pronotum (Fig. 49D): Margins of metazone with numerous small, sharp tubercles. Prothoracic Legs: Posterior prothoracic femoral genicular spine tiny, barely present, originating at the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third longer. Anterior surface of prothoracic coxae with a black medial mark in the proximal half.
Wings: Forewings colored symmetrically. Hindwings smoky black with rusty coloration at the very base of the discoidal region, sometimes extending distally along anterior margin; the terminus of the discoidal region projecting slightly beyond the distal margin of anal region.
Abdomen: Widening from first segment until the beginning of the distal half (segments 5-6) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites with slight posterolateral tergal projections in the distal half of the abdomen. Supra-anal plate slightly transverse, tapering to a rounded terminus.
Etymology. A noun in the genitive case, L. algorei is named for Albert Arnold 'Al' Gore, Jr., former Vice President of the United States of America, for his environmental activism including his efforts to raise public awareness of global climate change. Diagnosis. Nearly identical to L. nubeculosa, the primary distinguishing characteristics being male genital features, distribution and the hindwing. It is one of the largest Liturgusa species with robust legs, highly contrasting mottled forewings and strong banding. A feature easily seen on most specimens that is capable of distinguishing the species from L. nubeculosa is that the discoidal region of the hindwing does not extend much beyond the distal margin of the anal region, which gives the wing a more truncate appearance compared to the hindwing of L. nubeculosa.
Pronotum (Fig. 49E): Elongate, but robust with a slightly defined supra-coxal bulge; dorsal surface with numerous small tubercles. Prozone elongate with lateral margins that are near parallel, tapering anteriorly; the margins smooth. Metazone with concave lateral margins, a slight bulge in the posterior half; margins with few small, blunt tubercles; posterior margin slightly emarginate medially; the anterior half of dorsal surface raised, the dorsal surface of the posterior half depressed. Mostly brown with black marks.
Prothoracic Legs: Femur elongate with a slightly concave dorsal margin; strongly defined pale to dark banding on posterior (external) surface; anterior (internal) surface with a thin, black band running medially from the base to terminus; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned just distal and medial to the most proximal posteroventral spine; pit is dark. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, 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 longer. Prothoracic coxae smooth; the anterior surface with a small, black band medially in the proximal half as well as a small black spot medially towards the distal terminus.
Meso-and Metathoracic Legs: Femora with strongly pronounced ventral (posterior) and dorsal (anterior) carina; posterior surface with medial carina in the distal two thirds. Mesotarsi with first segment at most as long as the remaining segments combined.
Wings: Forewings mottled with brown, green, and pale coloration; the costal region with pale and dark irregular banding. Forewings sometimes colored asymmetrically, one being mottled the other is rust colored with the mottled pattern still visible. Hindwings smoky black and brown, opaque; the terminus of the discoidal region not projecting much beyond the distal margin of anal region, giving the wing a truncate appearance.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; smooth. Tergites with small posterolateral tergal projections. Supra-anal plate transverse, tapering to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52H.1): The main body of ventral left sclerite (L4A) with a sharply pointed distal process (pda) resembling a curved tooth positioned laterally, the terminal margin of the L4A centrally rounded before a depression that leads to the pda. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) broad and tapering quickly to a point; the apical process (paa) broad, cylindrical and curved, the terminus evenly rounded. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded, membranous terminus with fine setae; the ventral plate (pia) strongly sclerotized, broadening proximally with strong curved grooves; the ventral process (pva) c-shaped, tapering to a point distally.
Head (Fig. 44F): Slightly transverse, juxta-ocular protuberances medium, the apex in the middle third; the vertex between the parietal sutures is slightly concave and broad; vertex well above the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc.
Pronotum (Fig. 49F): Elongate, but robust with a moderately defined supra-coxal bulge. Metazone with concave lateral margins, a slight bulge in the posterior half; margins with small, blunt tubercles; posterior margin slightly emarginate medially.
Prothoracic Legs: Femur with very long posteroventral spines. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is very deep, but pigmented pale. Prothoracic tibial posteroventral spines with the first (proximal) very small and the third through sixth of similar length, the second very long, nearly the same length as the terminal spine.
Meso-and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings broadened with a widened costal region, extending at most to the tip of the abdomen, but usually shorter; mottled with highly contrasting brown and pale coloration; the costal region with pale and dark irregular banding. Forewings colored symmetrically. Hindwings smoky black and brown, opaque, the anterior margin more pale.
Abdomen: Broad, widening from first segment until the beginning of the distal half (segment 4) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites in the posterior half with small posterolateral tergal projections. Supra-anal plate as long as wide, evenly rounded.
Etymology. A noun in the genitive case, L. cameroni is named for Stephen L. Cameron for his contributions to Mantodea sampling and his valuable collaboration.  Natural history. Males and female found in local abundance at the Madre Selva Biological Research Station in the Loreto Province, Peru. The species was living in sympatry with L. algorei on the same smooth bark, medium diameter trees. The species was easily collected during the day and often found at reachable heights in lower sections of the tree.
Diagnosis. Most similar to L. purus and L. algorei in body shape and slenderness, but far more green in coloration across head, pronotum, and prothoracic legs with highly contrasting banding across the body. In addition, the head is mostly pale, but the lower portion of the frons, the clypeus, mandibles and labrum are darkly colored. Forewings are browner than other two species with a pronounced green and dark banded costal region. Hindwings are darker than L. purus, but more rusty than L. algorei. In addition, the prothoracic femoral posterior genicular spine in females is elongate, but shorter than posteroventral spines, but relatively much longer than that seen in L. algorei.
Head (Fig. 44G): Transverse, juxta-ocular protuberances medium, the apex in the lateral third; the vertex between the parietal sutures is straight; vertex even with the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and protruding on a small cuticular mound, but almost flat. Lateral ocelli oriented outward, a few degrees off perpendicular. Upper margin of clypeus barely convex, lower margin straight. Antennae pale basally fading quickly to black. Broad black band extending straight over the medial carina of the frontal suture; lower region of frons with two black marks separated by a pale gap centrally; the clypeus brown on lateral and lower region; the mandibles and labrum dark brown; the vertex and juxta-ocular protuberances splotched with brown and green; the area adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 49G): Elongate with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone with lateral margins that are near parallel, tapering anteriorly; the margins smooth. Metazone with concave lateral margins, a slight bulge in the posterior half; margins with numerous small tubercles; posterior margin medially emarginate; the dorsal surface of the posterior half moderately depressed. Green coloration dominant, but a few strong black marks.
Prothoracic Legs: Femur elongate 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; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is pale. Poste- rior prothoracic femoral genicular spine at most a quarter the length of the posteroventral spines, originating just distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the second through sixth of similar length, the second and third are slightly longer than adjacent spines. Prothoracic coxae smooth; the anterior surface with a broad, black band medially in the proximal half as well as a black spot medially towards the distal terminus.
Wings: Forewings mottled with brown, green, and pale coloration; the costal region with green and dark irregular banding. Forewings colored asymmetrically, one being mottled the other is rust colored with the mottled pattern still visible. Hindwings with a red or rusty color in the discoidal region, darkening distally to black; the anal region rusty basally and fading to a smoky grey and translucent medially before becoming black along the distal margin; the terminus of the discoidal region projecting well beyond the distal margin of anal region, the wings appearing elongate.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; smooth, black and green colored dorsal surface. Supra-anal plate transverse, tapering to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52I.1): The main body of ventral left sclerite (L4A) with a dull or sharply pointed, sickle shaped distal process (pda) positioned laterally, the terminal margin of the L4A centrally rounded before a depression that leads to the pda. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) straight, tapering into a needle-like process; the apical process (paa) broad, cylindrical and curved, the terminus a blunt and rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus, both margins sclerotized with a broad central membranous gap; the ventral plate (pia) strongly sclerotized and broad, with strong and numerous, curved grooves; the ventral process (pva) c-shaped, tapering to a point distally.
Pronotum (Fig. 49H): Prozone with lateral margins that are parallel, tapering anteriorly; the margins smooth. Metazone with concave lateral margins, a barely visible bulge in the posterior half; margins with small tubercles medially. Highly contrasting green and black coloration.
Prothoracic Legs: Femoral pit is black. Posterior prothoracic femoral genicular spine smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the fourth through sixth of similar length, the second and third are much longer.
Wings: Forewings colored symmetrically or asymmetrically, one being rust colored. The anal region of hindwings rusty basally and fading to a smoky grey and translucent medially before becoming black along the distal margin; the terminus of the discoidal region projecting beyond the distal margin of anal region.
Abdomen: Widening from first segment until the beginning of the distal half (segments 5-6) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Supra-anal plate as long as wide, evenly rounded.
Etymology. A noun in the genitive case, L. krattorum is named for Chris and Martin Kratt, hosts and creators of Kratts' Creatures and Wild Kratts, among other television shows, which provide children with entertaining programming focused on animal biology presented with accurate information.
Liturgusa maroni sp. n. http://zoobank.org/3406E298-A77C-4C78-A89B-039B6A80C540 http://species-id.net/wiki/Liturgusa_maroni Diagnosis. A medium size species known only from female specimens from French Guiana. Most similar to L. nubeculosa, but much smaller and with blunt and shortened hindwings. The pronotum is moderately elongate and females have a broad abdomen. Forewings slightly shortened and obviously rounded, extending barely to the tip of the abdomen.
Head (Fig. 45B): Slightly transverse, juxta-ocular protuberances large, the apex in the middle third; the vertex between the parietal sutures is slightly concave; vertex above dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward. Upper margin of clypeus barely convex, lower margin concave. Black markings surrounding frontal suture, the lower forming a point medially; lateral margins of frons with two, small black marks; the clypeus pale; the mandibles dark brown; the labrum mostly pale; the vertex and juxta-ocular protuberances splotched with dark brown marks; the area adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 49I): Elongate with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone with lateral margins that broaden slightly anterior to the supra-coxal sulcus before tapering anteriorly to a rounded terminus. Metazone with concave lateral margins, becoming nearly parallel in the posterior half with a slight bulge prior to narrowing towards the rounded posterior terminus; margins with small tubercles; posterior margin medially emarginate; the dorsal surface of the posterior half moderately depressed. The two posterior bulges on each side of the medial line are pronounced. Mostly brown with few black markings.
Prothoracic Legs: Femur elongate 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; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is pale. Posterior prothoracic femoral genicular spine smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) and fourth through sixth approximately the same size, the third being longer and the second very long, being nearly as long as the terminal spine. Prothoracic coxae smooth; the anterior surface with a broad, black band medially in the proximal half as well as a black spot medially towards the distal terminus.

Meso-and Metathoracic Legs:
Femora with ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment slightly shorter or the same length as the remaining segments combined.
Wings: Forewings mottled with brown, black, and pale coloration that is highly contrasting with large regions of lighter colored splotches; the costal region widened slightly with irregularly spaced black and pale banding; the terminus rounded and blunt, the overall shape appearing like a paddle, not extending to the tip of the abdomen. Forewings colored symmetrically. Hindwings smoky grey and translucent, the costal region more pale; the terminus of the discoidal region not projecting beyond the distal margin of anal region, the wing appearing truncate.
Abdomen: Broad, widening from first segment until the beginning of the distal half (segment 4) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites in the posterior half with small posterolateral tergal projections. Supra-anal plate triangular with rounded margins and a rounded point.
Etymology. A noun in apposition, L. maroni is named for the Maroni River near the type locality of Saint Laurent du Maroni on the border of French Guiana and Suriname.
In 1954, James Rehn synonymized L. peruviana Giglio-Tos, 1914, with its senior synonym, L. nubeculosa Gerstaecker, 1889, but his action appears to have been overlooked. Here again these two species are synonymized, but credit for first recognizing this synonymy goes to Rehn. Natural history. Liturgusa nubeculosa is one of the largest species of Neotropical bark mantis. Males and females are extremely difficult to catch during the day and have been primarily found on large diameter, smooth bark trees. Once they spot a threat, they run rapidly up the tree and out of reach, often circling to the back side, which is typical of the genus. Males were collected far more often than females on a recent trip to the Loreto province in northern Peru, but the opposite was true at a location in Bolivia (see material examined). It is not clear whether there are distinct sex ratios in certain locations or this sex bias was coincidence of habitat utilization.
Diagnosis. Nearly identical to L. cameroni, the primary distinguishing characteristics being male genital features, distribution and the hindwing. It is one of the largest Liturgusa species with robust legs, highly contrasting mottled forewings and strong banding. A feature easily seen on most specimens that is capable of distinguishing the species from L. cameroni is that the discoidal region of the hindwing extends beyond the distal margin of the anal region, which gives the wing a slightly more elongate appearance compared to the hindwing of L. cameroni.
Head (Fig. 45C): Transverse, juxta-ocular protuberances small, the apex in the lateral third; the vertex between the parietal sutures is straight; vertex even or slightly below the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward. The frons transverse; lower region entirely black. Clypeus transverse; upper margin convex, lower margin concave. Antennae pale basally, fading to black within the proximal ten percent. A moderate black band that fades medially extending across the frontal suture from eye to eye; clypeus mostly pale with brown markings in the upper lateral corners; mandibles pale medially and brown laterally; the vertex and juxta-ocular protuberances speckled with dense black markings; the area adjacent to lateral ocelli black. Palpi are pale.
Pronotum (Fig. 49K): Elongate, but robust with a slightly defined supra-coxal bulge; dorsal surface with numerous small tubercles. Prozone elongate with slightly widening lateral margins before tapering anteriorly; the margins smooth. Metazone with concave lateral margins, a slight bulge in the posterior half; margins with small, blunt tubercles; posterior margin slightly emarginate medially; the anterior half of dorsal surface raised; the dorsal surface of the posterior half depressed. Mostly brown with black marks.
Prothoracic Legs: Femur elongate, but robust (thick) 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; the ventral surface pale. Posterior surface of femur with numerous tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is dark. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, 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 longer. Prothoracic coxae smooth; the anterior surface with a small, black band medially in the proximal half as well as a small black spot medially towards the distal terminus.
Meso-and Metathoracic Legs: Femora with strongly pronounced ventral (posterior) and dorsal (anterior) carina. Mesotarsi with first segment shorter than the remaining segments combined.
Wings: Forewings mottled with black, pale, brown and grey coloration that is highly contrasting, exhibiting large grey regions surrounded by black markings; the costal region widened with pale and dark irregular banding. Forewings colored asymmetrically, one being mottled as described the other is darkened significantly with a black or rust tone, the mottled pattern still visible. Hindwings smoky black, mostly translucent; the discoidal region with black, opaque pigmentation basally and in the anterior two thirds extending from the base to the distal terminus, continuing into the costal region; the terminus of the discoidal region projecting beyond the distal margin of anal region, giving the wing an elongate appearance.
Abdomen: Elongate, tubular with slight widening before the posterior narrowing; smooth and black coloration dorsally. Tergites without posterolateral projections. Supra-anal plate transverse, half as long as wide, tapering to a rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52J.1-J.2): The main body of ventral left sclerite (L4A) with a short, triangular distal process (pda) positioned lateral to the central axis, the outer margin straight, heavily sclerotized and serrated from the terminus of the pda extending proximally one quarter the length of L4A. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) large and broad, but short, forming a sharply pointed process with a convex medial margin (closest to the paa) leading to the terminus and a concave outer margin (opposite the paa), the entire structure resembling a broad sickle; the apical process (paa) broad basally, quickly narrowing to a heavily sclerotized, rounded terminus. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a broadly rounded, membranous terminus, the outer margin sclerotized; the ventral plate (pia) strongly sclerotized, broad proximally with strong curved grooves; the ventral process (pva) enlarged and c-shaped.
Redescription. Female. (Figs 1C   Head (Fig. 45D): As broad as long, juxta-ocular protuberances very large, the apex in the middle third; the vertex between the parietal sutures is slightly concave; vertex well above the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc, more angular medially than in male. Ocelli small and protruding on a contiguous carina connecting all three ocelli; lateral ocelli oriented outward, the central ocelli almost vertical. The frons transverse, lower region below the carina black. Clypeus slightly transverse. Frontal suture without a defined black band as described in males; clypeus pale; mandibles pale; the vertex and juxta-ocular protuberances speckled with dense black markings; the area adjacent to lateral ocelli black. Palpi are pale.
Prothoracic Legs: Prothoracic tibial posteroventral spines with the first (proximal) smallest and the third through sixth of similar length (the third slightly longer than the other three), the second much longer. Posterior prothoracic femoral genicular spine tiny compared to posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic coxae smooth; the anterior surface with a small, black band medially in the proximal half, but no black mark in near the distal terminus.
Meso-and Metathoracic Legs: As described for males. Wings: Forewings colored symmetrically. The terminus of the discoidal region of the hindwing projecting slightly beyond the distal margin of anal region, giving the wing a moderately elongate appearance.
Abdomen: Broad, widening from first segment until the beginning of the distal half (segment 4) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites with or without small posterolateral tergal projections in the posterior half of the abdomen. Supra-anal plate slightly transverse, rounded. Diagnosis. Brown and light green with a slender appearance. Forewings asymmetrically colored and hindwings with the discoidal region mostly orange or rust colored. Most similar to L. krattorum and L. algorei, but with a paler head and a less contrasting banding pattern. The prothoracic femora are more brown across the posterior surface compared to the other two species. In addition, L. purus has a posterior prothoracic femoral genicular spine that originates well proximal to the beginning of the genicular lobe, which is unique among L. krattorum, L. algorei, and other Cursor Group species.
Head (Fig. 45A): Transverse, juxta-ocular protuberances large, the apex in the lateral third; the vertex between the parietal sutures is straight; vertex even with the dorsal margin of the eyes. Frontal suture with a medial carina forming a continuous arc. Ocelli small and protruding on a small cuticular mound, but almost flat. Lateral ocelli oriented outward, a few degrees off perpendicular. Upper margin of clypeus convex, lower margin straight. Antennae pale basally fading quickly to brown. Broad black band extending straight over the medial carina of the frontal suture; lower region of frons dark brown with two pale lateral spots; vertex and juxta-ocular protuberances mostly brown; the area around ocelli pale, the lower region of clypeus, the mandibles, and the labrum are mostly brown. Palpi are pale.
Pronotum (Fig. 49J): Elongate with a moderately defined supra-coxal bulge; dorsal surface entirely smooth. Prozone with lateral margins that are near parallel, tapering anteriorly; the margins smooth. Metazone with concave lateral margins; margins with numerous small tubercles; posterior margin medially emarginate; the dorsal surface of the posterior half slightly depressed.
Prothoracic Legs: Femur elongate with a nearly straight dorsal margin; less defined pale to dark banding on posterior (external) surface, more brown on the ventral margin with strongly brown regions between posteroventral spines; anterior (internal) surface with a black band running medially from the base to terminus; the ventral surface pale. Posterior surface of femur with few tubercles. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is pale. Posterior prothoracic femoral genicular spine half the length as the posteroventral spines, originating well proximal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the second through sixth of similar length, the second and third are slightly longer. Prothoracic coxae smooth; the anterior surface with a broad, diagonal black band in the proximal half.
Wings: Forewings mottled with brown, green, and pale coloration; the costal region with light to dark irregular banding. Forewings colored asymmetrically, one being mottled, the other is rust colored with the mottled pattern still visible. Hindwings with a red or rusty color in the discoidal region, darkening distally; the anal region a smoky grey and translucent; the terminus of the discoidal region projecting well beyond the distal margin of anal region, the wings appearing elongate.
Abdomen: Elongate, tubular with slight widening before posterior narrowing; smooth, reddish dorsal surface. Supra-anal plate transverse, tapering to a broadly rounded terminus. Subgenital plate irregularly rounded and without styli.
Genital Complex (Fig. 52K.1): The main body of ventral left sclerite (L4A) with a bulging distal process (pda) positioned laterally (45 degrees from central axis) and projecting like a blunt knob. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) broadly tapering to a short, sharp point that is well sclerotized and oriented approximately 25 degrees from the central axis; the apical process (paa) broad, cylindrical and curved, the terminus a tapered and rounded end. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus; the ventral plate (pia) strongly sclerotized, slightly widened proximally with curved grooves; the ventral process (pva) c-shaped, tapering to a point.
Female. (Fig. 29B) A single specimen is known, but has degraded due to poor preservation. Measurements are impossible, but some aspects of the description are possible and those included are presented in full as to not confuse character states that may or may not match with males.
Head: Frontal suture with a medial carina forming a continuous arc. Ocelli small and almost flat on the surface. Upper margin of clypeus convex, lower margin straight. Antennae pale basally fading quickly to brown. Palpi are pale.
Prothoracic Legs: Femur with less defined pale to dark banding on posterior (external) surface, more brown on the ventral margin with strongly brown regions between posteroventral spines; anterior (internal) surface with a black band running medially from the base to terminus, a broad mark medially; the ventral surface pale. A well developed femoral pit to accommodate terminal posteroventral tibial spine positioned medially to the proximal two posteroventral spines and in line with the distal most discoidal spine; pit is pale. Prothoracic tibial posteroventral spines with the first (proximal) smallest and the second through sixth of similar length, the second and third are slightly longer. Prothoracic coxae smooth; the anterior surface with a broad, diagonal black band in the proximal half.
Meso-and Metathoracic Legs: Mesotarsi absent, metatarsi with first segment much longer than remaining segments combined.

Type species. Mantis annulipes Audinet Serville, 1838
Description. Body: The overall coloration of all Fuga species varies with a mottled or camouflage pattern that incorporates black, brown, pale tan, white or grey, and sometimes shades of green. The mottled patterns can be diffuse or highly contrasting with whitish regions abutting black spots or splotches. All species are dorsoventrally flattened with disproportionately long legs in comparison to body length.
Head: Transverse with large, rounded eyes projecting outside the profile of the head both laterally and anteriorly (the anterior margin of the eyes anterior to the central surface of the head). Juxta-ocular protuberances present to varying degrees within males, but always well developed in females. The vertex between the parietal sutures is either straight or concave. Frontal suture with a faint medial carina. Ocelli small in males protruding slightly on a cuticular mound; reduced in females and laying more flat on the surface. Central ocellus oriented anteriorly and lateral ocelli oriented outward, perpendicular to the central axis of the head or at most a few degrees off perpendicular. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus convex, lower margin straight; a transverse ridge medially; lateral margins tapering, widest at the upper margin. Labrum with minimal sculpting and a rounded terminus. Antennae filiform and with rare setae, pale or dark or a combination of both, never banded. Varying levels of black markings across the anterior surface of head that can include a transverse band or spots on the lower part of the frons, markings around the ocelli and the vertex, and markings on the clypeus, labrum and mandibles. Palpi are pale or with a darkened terminus.
Pronotum: Slightly elongate (pronotum shape measure 0.36) to squat (pronotum shape measure 0.49) with a moderately defined supra-coxal bulge; dorsal surface smooth or at most with dispersed, fine tubercles. Prozone with lateral margins that are parallel before tapering anteriorly. Metazone with concave lateral margins, always with a middle region that is narrower than the supra-coxal bulge and the posterior end. Coloration highly variable with pale and black markings. Supra-coxal sulcus strongly defined; posterior margin straight or barely medially emarginate.
Prothoracic legs: Femoral spine count of male and female: anteroventral 14-17, posteroventral 4, discoidal 4. Femur robust with a straight or slightly concave dorsal margin; anteroventral and posteroventral (internal and external, respectively) spines well developed; line of small tubercles running medially of the posteroventral spines. A continuous carina running from distal terminus of femur along dorsal margin to the base, circling the posterior surface of the proximal end and running along the ventral margin at the base of the posteroventral spines. Pale to dark banding on posterior (external) surface of femur; anterior (internal) surface entirely black or pale with varying patterns of black markings. Posterior (external) surface of femur smooth or with few tubercles. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned between the most proximal posteroventral spine and the most distal discoidal spine; pit is colored black or pale. Prothoracic tibial spine count of male and female: anteroventral 9-10, posteroventral 8. Prothoracic tibial spines robust; the posteroventral spines with the first and second most proximal and fifth through seventh shorter than the longer proximal third, fourth, and terminal spines; the anteroventral spines longest at distal end and shortening proximally, but the sixth and seventh spines from the distal terminal spine longer than adjacent spines. Tarsi banded with pale and dark coloration. Prothoracic coxae smooth with no or a few very minor tubercles or setae along dorsal margin; black markings vary across species.
Meso-and metathoracic legs: Long and slender with pale to dark banding on the femur and tibia; posterior (upper) surface of femora smooth. Femora with ventral (posterior) carina, some species being more pronounced than others; dorsal (anterior) carina less pronounced, but visible. Tibia long and rounded with well developed terminal spurs. Mesotarsi with first segment as long or shorted than the remaining segments combined. Metatarsi with first segment equal to or longer than remaining segments combined.
Wings: Developed in males and females. Forewings mottled with contrasting regions of brown, green, and pale tan, and sometimes dark black; the costal region wide relative to the wing length, the width between 4-8% the total wing length, always with light -dark regular banding. The forewings may be colored asymmetrically, one being mottled as described above while the other is either dark rust or blackened with the mottled pattern still slightly visible (darker wing typically folded under the mottled wing). Hindwings opaque and smoky; the distal terminus of the discoidal region darker than the rest; the costal region can be much darker or paler than the discoidal region depending on species; the terminus of the discoidal region projecting well beyond the distal margin of anal region, making the wing appear pointed or elongate.
Abdomen: Males and females with gradually widening abdomen from first segment until the beginning of the distal half (segments 5-7) when the lateral margins narrow to the terminus, the middle being the broadest region. Posterolateral corners of tergites simple, without projections. Cerci cylindrical, long and setose, tapering to a point. Supra-anal plate long (females) or transverse (males), with a rounded or more pointed terminus. Subgenital plate of male with rounded, slightly irregular terminus; without styli.
Male genital complex: The main body of ventral left sclerite (L4A) with a prominent and curved distal process (pda). The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) well sclerotized with a blunt, rounded terminus; the apical process (paa) cylindrical and curved, terminating with a rounded end; with or without a large membranous lobe originating between the apofisis falloid (afa) and the apical process (paa), if present then with or without robust setae. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus and is mostly membranous with setae; the ventral plate (pia) strongly sclerotized and short, but with a smooth surface; the ventral process (pva) strongly sclerotized and curved.
When describing Mantis annulipes in 1838 Audinet Serville provided a relatively lengthy description, but was ambiguous when referencing the locality of the specimen or specimens by only listing "Of Brazil and of Cayenne". He did reference that the description was based on the female form, but it is unknown if there were one or multiple specimens examined; the distribution information presented suggests multiples. The specimens are presumed to come from the collection of MM. Dejean et Viard, but the current location of Audinet Serville's specimens is not known. The catalog of Reinhard Ehrmann listed the holotype female as deposited in the Muséum national d'Histoire naturelle, Paris, and paratypes deposited in the Ernst-Moritz-Arndt-Universität Greifswald, Germany. The specimens from EMAU were located and examined, but the holotype in MNHN was not located. A search did not reveal its presence in the collection historically. It is not truly known whether the two EMAU specimens are those examined by Audinet Serville, but they date from the correct period and are from locations in Brazil. However, since a holotype was never designated by Audinet Serville and no subsequent fixation was uncovered, the specimens are treated as syntypes. Therefore, to increase taxonomic stability within the species, under Article 74.1 of the International Code of Zoological Nomenclature a the female from Bahia, Brazil deposited in EMAU has been selected to become the unique bearer of the name of the nominal species-group Fuga annulipes (lectotype). The additional specimen also deposited in EMAU is selected as a paralectotype under Article 74.1.3 of The Code.
The types of three species, Hagiomantis parva Piza, 1966, Liturgusa sinvalnetoi Piza, 1982, and Liturgusa parva Giglio-Tos, 1914, were examined and compared with the original description of Mantis annulipes and the two specimens from EMAU and were found to be conspecific. These three new synonymies are likely the result of the historical confusion surrounding the distribution and characteristics of Fuga annulipes.
Diagnosis. A small species, males being darkly colored with highly contrasting grey and black on forewings. Females also with a darker coloration, but most noticeable is the less elongate pronotum compared to the other two species in the genus. Their squat appearance should be obvious when observed in the field as they are the only species known in eastern Brazil with such short pronota.
Head (Fig. 45E): Juxta-ocular protuberances small, the middle being the most pronounced; the vertex between the parietal sutures is slightly concave; vertex lower than the dorsal margin of the eyes. Frontal suture with a curved carina forming a continuous arc. Ocelli small with the central ocellus oriented anteriorly while the lateral are oriented slightly off 90 degrees from the central axis of the head; protruding slightly on cuticular mounds. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus slightly convex, lower margin slightly concave; a central protruding carina; the lateral margins tapering, widest at the upper margin. Antennae pale proximally and fading to brown distally. Varying levels of black markings across the anterior surface of head; the vertex, juxta-ocular protuberances, frons, mandibles, and labrum with black and pale mottling, but giving a darker appearance. Palpi are darkened terminally.
Pronotum (Fig. 50A): Shortened, but not squat with a slightly defined supra-coxal bulge; dorsal surface mostly smooth with a few tubercles present; prozone with parallel lateral margins prior to a rounded anterior margin; metazone with concave lateral margins, smooth or at most with tiny tubercles associated with setae; posterior margin of the metazone rounded with a slight medial emargination; the dorsal surface of the posterior half of the metazone slightly depressed. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens.
Prothoracic Legs: Femur robust with a straight dorsal margin. Brown to dark banding on posterior (outer) surface of femur with less than 10 tubercles; anterior (internal) surface amber or pale with one black spot in the groove for the tibial spur; ventral surface amber or pale. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned medial to the most proximal posteroventral spine; pit is black. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial posteroventral spines with the first and second (proximal) short with the fourth through seventh of similar length, the third and terminal spines much longer; the anteroventral spines longest at distal end and shortening proximally. Posterior surface of the prothoracic tibiae smooth and banded, but dark brown; anterior surface amber, the ventral surface amber. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a proximal (near center) and a distal black marking.
Meso-and Metathoracic Legs: Femora with a pronounced dorsal and ventral carina; posterior (upper) surface with two carinae, one running nearly the entire length, fading proximally and positioned ventrally to the dorsal carina, the other in the distal third and positioned parallel and between the previous and the ventral carina. Mesotarsi with first segment as long as the remaining segments combined. Metatarsi with first segment slightly longer than remaining segments combined.
Wings: The same length or slightly longer than the abdomen. Forewings mottled with contrasting regions of black and pale white or grey; the costal region alternating from pale to dark its entire length, mostly brown proximally; the costal region slightly widened. The forewings may be colored asymmetrically, one being mottled as described above while the other is much darker and usually folded under the other. Hindwings opaque and smoky with near black veins; the terminus of the discoidal region truncate and not projecting beyond the distal margin of anal region, giving a stubby appearance.
Head (Fig. 45F): Juxta-ocular protuberances moderate size, the middle being the most pronounced; the vertex between the parietal sutures is slightly concave; vertex higher than the dorsal margin of the eyes. Ocelli protruding slightly on a carina connecting all three and extending laterally. Varying levels of black markings across the anterior surface of head; the vertex, juxta-ocular protuberances, frons, mandibles, and labrum with fine black speckling over a largely pale coloration, giving a lighter appearance than males. Palpi are darkened brown terminally.
Pronotum (Fig. 50B): Shortened, but not squat with a slightly defined supra-coxal bulge; dorsal surface mostly smooth, but some tubercles are present; prozone with parallel lateral margins prior to a rounded anterior margin; metazone with concave lateral margins, with small tubercles in the posterior two thirds; posterior margin of the metazone rounded with no medial emargination.
Prothoracic Legs: Femur robust with a nearly straight dorsal margin; anteroventral spines black, posteroventral spines pale basally with a black terminus. Prothoracic tibial posteroventral spines with the first and second most proximal and fifth through seventh shorter than the proximal third, fourth and terminal spines; the anteroventral spines longest at distal end and shortening proximally, but the sixth and seventh from the distal end longer than adjacent spines. Posterior surface of the prothoracic tibiae smooth and banded; anterior surface pale, the ventral surface pale. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a small proximal (near center) and a small distal black marking.
Meso-and Metathoracic Legs: Femora with a pronounced dorsal and ventral (posterior) carina. Mesotarsi with first segment as long as or shorter than the remaining segments combined. Metatarsi with first segment slightly longer than remaining segments combined.
Wings: Costal region of hindwings and the anterior margin of the discoidal region more pale than the rest.
Abdomen: Broad and smooth, widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites with small posterolateral projections on the sixth and seventh segments. Supra-anal plate longer than wide, broadly rounded. Taxonomic history. Described in 1965 by Piza as a species of Hagiomantis, the species has since received no attention taxonomically.
Diagnosis. Medium size species, mottling pattern highly contrasting with light and dark browns, no white, black, or green included. Costal region of the forewing one of the widest in the genus relative to the wing length, banded with regular alternating patterning between pale and dark brown.
Pronotum (Fig. 50C): Slightly elongate with a slightly defined supra-coxal bulge; dorsal surface mostly smooth, but with a few very small tubercles; prozone with parallel lateral margins prior to a rounded anterior margin. Metazone with concave lateral margins with a slight medial bulge, exhibiting small denticles in the posterior two thirds; posterior margin of the metazone with angled margins leading to a straight or slightly concave posterior margin, no noticeable medial emargination; the dorsal surface of the posterior half of the metazone depressed. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens.
Prothoracic Legs: Femur robust with a straight or slightly concave dorsal margin; anteroventral spines black, posteroventral spines pale basally with black terminus. Black and pale mottling on posterior (external) surface of femur with less than 10 tubercles; anterior (internal) surface entirely black; ventral surface entirely black. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned medial to and slightly proximal to the most proximal posteroventral spine; pit is black. Posterior prothoracic femoral genicular spine smaller than posteroventral spines, originating at the beginning of the genicular lobe. Prothoracic posteroventral tibial spines with the first and second most proximal and fourth through seventh shorter than the proximal third and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Posterior surface of the prothoracic tibiae smooth and dark; anterior surface black, the ventral surface brown. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a medial black stripe.
Wings: Longer than the abdomen. Forewings mottled with contrasting regions of light and dark brown; the costal region alternating regularly from pale to dark its entire length; the costal region slightly widened. The forewings may be colored asymmetrically, one being mottled as described above while the other is much darker.
Hindwings opaque and smoky with near black veins; the costal region and the anterior margin of the discoidal region pigmented with a darker reddish or rust coloration; the terminus of the discoidal region projecting well beyond the distal margin of anal region, giving an elongate appearance.
Genital Complex (Fig. 53B.1-B.2): The main body of ventral left sclerite (L4A) with a prominent, robust, curved distal process (pda) that tapers quickly to a point, the entire structure curving laterally and back onto the L4A. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) small and curved, terminating with a blunt end; the apical process (paa) cylindrical and curved, terminating in a rounded end, slightly bulbous; a large, long membranous lobe originating between the apofisis falloid (afa) and the apical process (paa). Right phallomere (R1) too damaged for accurate description.
Head (Fig. 45H): Juxta-ocular protuberances pronounced, the apex in the middle; the vertex between the parietal sutures is straight or slightly concave; vertex higher than the dorsal margin of the eyes. Frontal suture with a curved carina forming a continuous arc. Antennae pale proximally and fading to black or dark brown distally. Varying levels of black markings across the anterior surface of head, variable within the species; the vertex and juxta-ocular protuberances dark brown; two black spots medial to the parietal sutures. Terminus of palpi are darkened.
Pronotum (Fig. 50D): Slightly elongate with a somewhat defined supra-coxal bulge; dorsal surface mostly smooth, but a few very small tubercles are present; prozone with parallel or divergent lateral margins prior to a rounded anterior margin; metazone with concave lateral margins with a medial bulge, exhibiting sparse small denticles in the posterior two thirds; posterior margin of the metazone rounded irregularly with a slight medial emargination. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens.
Prothoracic Legs: Black and pale banding on posterior surface of femur with less than 10 tubercles; anterior (inner) surface with a proximal black stripe as well as a distal black stripe medially; ventral surface with a black mark distally and a black mark just medial to second most proximal posteroventral spine. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating just distal to the beginning of the genicular lobe. Prothoracic posteroventral tibial spines with the first and second most proximal and fifth through seventh shorter than the proximal third, fourth and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Posterior surface of the prothoracic tibiae smooth with few black marks; anterior surface pale with black marks on dorsal margin, the ventral surface pale. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with two black markings of varying size, one in the proximal half and one in the distal half.
Meso-and Metathoracic Legs: Mesotarsi with first segment as long as the remaining segments combined. Metatarsi with first segment slightly longer than remaining segments combined.
Wings: Fully developed, longer than the abdomen. Forewings mottled with contrasting regions of light and dark brown; the costal region alternating regularly from pale to dark its entire length; the costal region widened, the widest part being in the middle of the wing. The forewings have not been observed to be asymmetrically colored.
Abdomen: Broad and smooth, widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites without posterolateral tergal projections. Cerci cylindrical, long and setose, tapering to a point. Supra-anal plate as long as wide, rounded. Diagnosis. Small species with a strongly defined darkened band running diagonally across the medial section of the forewing; forewings can be asymmetrically colored, one being darker than the other.
Head (Fig. 46A): Juxta-ocular protuberances pronounced, the apex in the middle; the vertex between the parietal sutures is straight; vertex slightly lower than the dorsal margin of the eyes. Frontal suture with a curved carina forming a continuous arc. Ocelli small with the central ocellus oriented anteriorly while the lateral are oriented slightly off 90 degrees from the central axis of the head; protruding slightly on cuticular mounds. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus slightly convex, lower margin slightly concave; a central protruding carina strongly defined; the lateral margins tapering, widest at the upper margin. Antennae pale proximally and fading to black distally. Varying levels of black markings across the anterior surface of head, variable within the species; the vertex and juxta-ocular protuberances dark brown. Palpi are pale.
Pronotum (Fig. 50E): Slightly elongate with a slightly defined supra-coxal bulge; dorsal surface smooth, but a few tubercles are present; prozone with parallel lateral margins prior to a rounded anterior margin; metazone with concave lateral margins, exhibiting small denticles in the posterior half; posterior margin of the metazone rounded and without a medial emargination; the dorsal surface of the posterior third of the metazone slightly depressed. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens. Prothoracic Legs: Femur robust with a straight dorsal margin; anteroventral spines black, posteroventral spines pale basally with black terminus. Pale to dark banding on posterior (outer) surface of femur with less than 10 tubercles; anterior (inner) surface black; ventral surface black. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned medial to and slightly proximal to the most proximal posteroventral spine; pit is black. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating distal to the genicular lobe. Prothoracic posteroventral tibial spines with the first and second most proximal and fifth through seventh shorter than the proximal third, fourth and terminal spines; the anteroventral spines longest at distal end and shortening proximally, but the sixth and seventh from the distal end longer than adjacent spines. Posterior surface of the prothoracic tibiae smooth and banded, but dark brown; anterior surface black; the ventral surface amber. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a proximal and a distal black marking.
Meso-and Metathoracic Legs: Femora with pronounced dorsal and ventral carinae. Mesotarsi with first segment as long as the remaining segments combined. Metatarsi with first segment longer than remaining segments combined.
Wings: Fully developed, the same length or slightly longer than the abdomen. Forewings mottled with contrasting regions of brown, white, and black; the costal region alternating regularly from pale to dark its entire length; the costal region slightly widened. The forewings may be colored asymmetrically, one being mottled as described above while the other is much darker and usually folded under the other. Hindwings opaque and smoky with near black veins; the costal region and the anterior margin of the discoidal region pigmented with a reddish or rust coloration; the terminus of the discoidal region projecting well beyond the distal margin of anal region, giving an elongate appearance.
Abdomen: Elongate, tubular, and smooth. Tergites with very small posterolateral projections in the distal half of the abdomen. Supra-anal plate transverse, evenly rounded. Subgenital plate rounded and without styli.
Genital Complex (Fig. 53C.1): The main body of ventral left sclerite (L4A) with a prominent, robust, curved distal process (pda) that tapers to a point, the entire structure curving laterally and back onto the L4A. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) well sclerotized with a blunt, rounded terminus, the tip broader than medial region; the apical process (paa) cylindrical and curved, terminating in a rounded end; a large, bifid and membranous lobe originating between the apofisis falloid (afa) and the apical process (paa), with robust setae along a lateral margin as well as emerging from the terminus of each sub-lobe. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a rounded terminus and is mostly membranous with setae; the ventral plate (pia) strongly sclerotized and short, but with a smooth surface; the ventral process (pva) strongly sclerotized and curved, terminating with a rounded, blunt end.
Head (Fig. 46B): Vertex slightly above the dorsal margin of the eyes. Pronotum (Fig. 50F): Metazone with concave lateral margins, exhibiting small denticles in the posterior two thirds; the dorsal surface of the posterior half of the metazone depressed.
Prothoracic Legs: Femur robust with a slightly concave dorsal margin distally, convex proximally; line of larger, pointed tubercles running medially of the posteroventral spines. Anterior (inner) surface of femur pale with proximal and dorso-distal black markings; ventral surface pale; femoral pit pale. Posterior prothoracic femoral genicular spine slightly smaller than posteroventral spines, originating just proximal to the genicular lobe. Posterior surface of the prothoracic tibiae smooth and banded; anterior surface mostly pale, the ventral surface pale. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a nearly continuous medial black marking running the entire length.
Meso-and Metathoracic Legs: Tibiae with a prominent ventral carina. Wings: The forewings rarely colored asymmetrically, if so then one being mottled while the other is much darker. Hindwings opaque and smoky with near black veins; the costal region and the anterior margin of the discoidal region more pale than the rest; the terminus of the discoidal region projecting well beyond the distal margin of anal region, giving an elongate appearance.
Abdomen: Broad and smooth, widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Tergites without posterolateral projections. Supra-anal plate as wide as long, tapering to a rounded point.
Etymology. A noun in the genitive case, L. grimaldii is named for David Grimaldi, whose work on fossil Mantodea added considerable knowledge to our understanding of the origins and evolution of the group.

Description. Habitus:
One of the larger Neotropical Liturgusini, Velox is long and slender with tapered forewings, an elongate pronotum and long, slender meso and metathoracic legs. Coloration is heavily mottled and most observed males and the single female have asymmetrical wing coloration, one wing being much darker and folded under the other.
Measurement Ranges: Monotypic genus, see V. wielandi for measurements. Head: Transverse with large, rounded eyes projecting outside the profile of the head both laterally and anteriorly (the anterior margin of the eyes anterior to the central surface of the head). Juxta-ocular protuberances present, the lateral third being the most pronounced. The vertex between the parietal sutures is straight or barely concave. Frontal suture with a faint curved carina forming a continuous arc. Ocelli small with the central ocellus oriented anteriorly while the lateral are oriented anterolaterally at a 45 degree angle from the central axis of the head; protruding slightly on cuticular mounds. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve, the middle forming an angle. Upper margin of clypeus straight, lower margin concave or straight; a central protruding ridge strongly defined; the lateral margins tapering, widest at the upper margin. Labrum with minimal sculpting and a rounded terminus; lateral margins widening ventrally. Antennae filiform and with rare setae, pale proximally and fading distally to light brown in males and dark brown in females. Varying levels of black markings across the anterior surface of head, variable within the species. Palpi are pale.
Pronotum: Elongate with a slightly defined supra-coxal bulge; dorsal surface with tubercles; prozone long with parallel lateral margins prior to a rounded anterior margin; metazone long with sweeping concave lateral margins in males and concave, but near parallel margins medially; margins exhibit small, occasional denticles; posterior margin of the metazone straight or broadly rounded, with a slight medial emargination. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens.
Prothoracic Legs: Femur robust with a straight dorsal margin distally, the proximal third slightly convex; anteroventral and posteroventral (internal and external, respectively) spines well developed; line of small, pointed tubercles running medially of the posteroventral spines; anteroventral spines black, posteroventral spines pale basally with black terminus. A strongly pronounced continuous carina running from distal terminus of femur along dorsal margin to the base, circling the external surface of the proximal end and running along the ventral margin at the base of the posteroventral spines. Pale to dark banding on posterior surface of femur with numerous tubercles; anterior (inner) surface amber colored or pale ventrally, a black strip medially, and pale along the dorsal margin; ventral surface between the anteroventral and posteroventral spines amber or pale colored. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned on the lateral margin, pushing the margin outward between the proximal two posteroventral spines; pit is pale. Prothoracic tibial spines robust; the posteroventral spines with the first and second most proximal and fourth through seventh shorter than the proximal third and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Posterior surface of the prothoracic tibiae smooth and banded with pale and dark coloration; anterior and ventral surface amber colored. Tarsi banded with pale and dark coloration. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface mostly black, but with varying pale marks.
Meso-and Metathoracic Legs: Long and slender with pale to dark banding on the femur and tibia. Femora with pronounced dorsal and ventral carinae; posterior (upper) surface smooth. Tibiae banded with alternating pale and dark regions with a pronounced ventral carina. Mesotarsi with first segment as long or shorter than the remaining segments combined. Metatarsi with first segment longer than remaining segments combined.
Wings: Fully developed, the same length or slightly longer than the abdomen. Forewings mottled with contrasting regions of brown, white, and black; the costal region alternating irregularly from pale to dark its entire length; the costal region widened. The forewings may be colored asymmetrically, one being mottled as described above while the other is much darker and usually folded under the other. Hindwings opaque and smoky with near black veins; the costal region and the distal tip of the discoidal region more opaque with black pigment; the terminus of the discoidal region projecting well beyond the distal margin of anal region giving the appearance of an elongate wing.
Abdomen: Elongate for males and broad for females. Tergites without posterolateral projections. Cerci cylindrical, long and setose, tapering to a point. Supra-anal plate highly to slightly transverse between the sexes. Subgenital plate of male elongate, tapering to the terminus with two prominent styli.
Genital Complex: The main body of ventral left sclerite (L4A) with a smooth, rounded terminus, the left side with a broad indentation. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) well sclerotized forming a smooth, rounded terminus in the shape of a broad, dull ninety degree hook; the apical process (paa) heavily sclerotized and curved, tapering to a smooth, rounded terminus that is heavily sclerotized. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a narrow, rounded terminus with short, dispersed setae; the ventral plate (pia) strongly sclerotized proximally, the surface rough and with broad curved grooves; the ventral process (pva) c-shaped, the distal end with rough, micro-toothed surface on the inside edge of the inward curve.
Ootheca. Unknown for the genus. Etymology. A substantivated adjective, the name is derived from the Latin noun "velox" meaning swift or rapid, an apt name for their amazing speed when running across tree trunks. Diagnosis. Large species with an elongate habitus. Distal terminus of the forewing tapered, giving a sharp appearance; often asymmetrically colored with one being much darker. Hindwings are elongate with the distal margin of the discoidal region projecting well beyond the anal region. Tubercles present across the pronotum, but meso and metafemora smooth on the posterior (upper) surface. The posteroventral spines of the prothoracic femora very long, the distal the largest of the four. Species similar in general appearance to H. mesopoda, but smaller and distributed in eastern Brazil.
Head (Fig. 46C): Juxta-ocular protuberances present, the lateral third being the most pronounced; the vertex between the parietal sutures is straight or barely concave; vertex just lower than the dorsal margin of the eyes. Frontal suture with a faint curved carina forming a continuous arc. Ocelli small with the central ocellus oriented anteriorly while the lateral are oriented anterolaterally at the 45 degree angle from the central axis of the head; protruding slightly on cuticular mounds. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve, the middle forming an angle. Upper margin of clypeus straight, lower margin concave; a central protruding ridge strongly defined; the lateral margins tapering, widest at the upper margin. Antennae pale proxi- mally and fading to light brown distally. Varying levels of black markings across the anterior surface of head. Palpi are pale.
Pronotum (Fig. 50G): Elongate with a slightly defined supra-coxal bulge; dorsal surface with tubercles; prozone long with parallel lateral margins prior to a rounded anterior margin; metazone long with sweeping concave lateral margins that exhibit small, occasional denticles; posterior margin of the metazone straight with a slight medial emargination; the dorsal surface of the posterior half of the metazone depressed. Colored with black and pale markings that vary across specimens.
Prothoracic Legs: Femur robust with a straight dorsal margin distally, the proximal third slightly convex. Pale to dark banding on posterior surface of femur with numerous tubercles; anterior (inner) surface amber colored ventrally, a black strip medially, and pale along the dorsal margin; ventral surface amber colored. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned on the lateral margin, pushing the margin outward between the proximal two posteroventral spines; pit is pale. Posterior prothoracic femoral genicular spine at most half the length of the posteroventral spines, originating proximal to the beginning of the genicular lobe. Prothoracic posteroventral tibial spines with the first and second most proximal and fourth through seventh shorter than the proximal third and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Posterior surface of the prothoracic tibiae smooth and banded with pale and dark coloration; anterior and ventral surface amber colored. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface mostly black with the ventral margin pale.
Meso-and Metathoracic Legs: Femora with pronounced dorsal and ventral carinae; posterior (upper) surface smooth. Tibiae with a pronounced ventral carina. Mesotarsi with first segment as long as the remaining segments combined. Metatarsi with first segment longer than remaining segments combined.
Wings: Fully developed, the same length or slightly longer than the abdomen. Forewings mottled with contrasting regions of brown, white, and black; the costal region alternating irregularly from pale to dark its entire length; the costal region widened. The forewings may be colored asymmetrically, one being mottled as described above while the other is much darker. Hindwings opaque and smoky with near black veins; the costal region and the distal tip of the discoidal region more opaque with black pigment; the terminus of the discoidal region projecting well beyond the distal margin of anal region, the wing appearing elongate.
Abdomen: Elongate, tubular, and smooth. Tergites without posterolateral projections. Supra-anal plate highly transverse with a blunt, straight terminus. Subgenital plate elongate, tapering to the terminus with two prominent styli.
Genital Complex (Fig. 53D.1): The main body of ventral left sclerite (L4A) with a smooth, rounded terminus, the left side with a broad indentation. The apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) well sclerotized forming a smooth, rounded terminus in the shape of a broad, dull ninety degree hook; the apical process (paa) heavily sclerotized and curved, tapering to a smooth, rounded terminus. The right dorsal phallomere (fda) of the first sclerite of right phallomere (R1) tapers to a narrow, rounded terminus with short, dispersed setae; the ventral plate (pia) strongly sclerotized proximally, the surface rough and with broad curved grooves; the ventral process (pva) c-shaped, the distal end with rough, micro-toothed surface on the inside edge of the inward curve.
Head (Fig. 46D): Vertex slightly higher than the dorsal margin of the eyes. Upper margin of clypeus straight, lower margin straight. Antennae pale proximally and fading to dark brown distally.
Pronotum (Fig. 50H): Elongate with a slightly defined supra-coxal bulge; dorsal surface with tubercles; prozone long with parallel lateral margins prior to a rounded anterior margin; metazone long with slightly concave lateral margins, nearly straight medially; lateral margin with very small, occasional denticles; posterior margin of the metazone broadly rounded with a slight medial emargination.
Prothoracic Legs: Anterior (inner) surface of femur pale ventrally, a black strip medially, and pale along the dorsal margin; ventral surface pale. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface with a medial black marking in the proximal half.
Meso-and Metathoracic Legs: Mesotarsi with first segment shorter than the remaining segments combined. Metatarsi with first segment longer than remaining segments combined.
Wings: As described for males. Abdomen: Broad, widening from first segment until the beginning of the distal half (segments 4-5) when the lateral margins narrow gradually to the terminus, the middle being the broadest region. Supra-anal plate slightly transverse with a dull, pointed terminus, the margins tapering distally.
Etymology. A noun in the genitive case, V. wielandi is named for Frank Wieland, whose contributions to Mantodea systematics, phylogeny and morphology have advanced the field considerably.
Measurement Ranges: Monotypic genus, see C. atricoxata for measurement ranges. Head: Transverse with large, rounded eyes projecting outside the profile of the head both laterally and anteriorly (the anterior margin of the eyes anterior to the central surface of the head). Juxta-ocular protuberances present, small in males and well developed in females. The vertex between the parietal sutures is straight. Frontal suture with a faint curved carina. Ocelli small in males and females with the central ocellus oriented anteriorly while the lateral are oriented anteriorly at a 45 degree angle from the central axis of the head; protruding on a cuticular mounds or on a prominent continuous curved carina. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus convex, lower margin straight; a central protruding carina; the lateral margins tapering, widest at the upper margin. Labrum with minimal sculpting and a rounded terminus. Antennae filiform and with rare setae, pale proximally and fading to black distally. Varying levels of black markings across the anterior surface of head, the vertex and juxta-ocular protuberances speckled. Palpi are pale.
Pronotum: Short and broad with a less defined supra-coxal bulge, the metazone being wide and the lateral margins of the metazone nearly parallel before tapering posteriorly; dorsal surface with tubercles of varying size and density, but obviously prominent. Broad prozone with lateral margins that taper anteriorly, the anterior margin rounded; a central depression medially on the dorsal surface. Metazone with lateral margins that are nearly parallel anteriorly, but taper dramatically to the narrowest point (the midpoint of the metazone) before becoming parallel anterior to the posterior terminus; the corners of the posterior margin rounded with the medial region straight; posterior region with two blunt protrusions on each side of the medial line. Supra-coxal sulcus strongly defined. Colored with black and pale speckling. The lateral margins of the pronotum slightly expanded to form a ledge.
Prothoracic Legs: Femoral spine count of males and females: anteroventral 15-16, posteroventral 4, discoidal 4. Femur robust with a slightly concave dorsal margin distally, the proximal half convex, larger in females; anteroventral and posteroventral (internal and external, respectively) spines well developed; line of small tubercles running medially of the posteroventral spines. A continuous carina running from distal terminus of femur along dorsal margin to the base, circling the posterior (external) surface of the proximal end and running along the ventral margin at the base of the posteroventral spines. Pale to dark banding on posterior surface of femur, sometimes not well defined and degenerating into pale and dark speckling ventrally; internal surface mostly black, but with pale regions dorsally; ventral surface black and pale. Posterior surface of femur smooth or with few tubercles. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned between the most proximal posteroventral spine of femur and the most distal discoidal spine, pit is colored black. Prothoracic tibial spine count of males and females: anteroventral 9-10, posteroventral 8. Prothoracic tibial spines robust; the posteroventral spines with the first and second most proximal and fifth through seventh shorter than the much longer proximal third, fourth, and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Prothoracic tibiae with a smooth posterior surface. Tarsi banded with pale and dark coloration. Prothoracic coxae smooth with no or a few very minor tubercles or setae along anterior margin; black markings vary.
Meso-and Metathoracic Legs: Long and slender with pale to dark banding on the femur and tibia. Femora with a pronounced dorsal and ventral (posterior) carina; posterior (upper) surface with two carina. Tibia widening from the proximal terminus and with multiple, faint carina. Mesotarsi with first segment shorter than remaining segments combined. Metatarsi with first segment the same length as the remaining segments combined.
Wings: Fully developed in males and females. Forewings mottled with contrasting regions of brown, white, green, and black; the proximal quarter dark, then fading dramatically to a mottled white with a darkened spot on and around the pterostigma; the distal quarter of the wing mottled irregularly with half greenish white and half brownish black; the costal vein alternating from pale to dark its entire length while the costal region is mostly pale with banding in the distal quarter, but can be banded throughout its length; the costal region wide relative to the wing length. The forewings may be colored asymmetrically, one being mottled as described above while the other is blackened. Hindwings opaque and smoky; the terminus of the discoidal region projecting to beyond the distal margin of anal region.
Abdomen: Males and females with widening abdomen from first segment until the beginning of the distal half (segments 5-6) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites with pointed posterolateral projections in the distal half of the abdomen of females. Cerci cylindrical, long and setose, tapering to a point. Supra-anal plate of female as broad as wide with a blunt terminus, large in size; of male transverse with blunt terminus. Subgenital plate of male with rounded, slightly irregular terminus; without styli.
Male Genital Complex: The only known species for the genus exhibits dextral genitalia ('reversed' genitalia in Balderson 1978 andHolwell andHerberstein (2010); defined as a genital complex in which the apical process (paa) of L4B is directed to Taxonomic history. First described by Max Beier, the species was referenced only a few times over the next 80 years and never treated in a comprehensive taxonomic study. However, Rehn (1935) suggested that the species did not really belong within Liturgusa based on its considerable differences in general appearance. Examining the male genitalia and redescribing the genus allowed description of numerous characters that distinguishes the species from all other Neotropical Liturgusini. More than one species likely exists, but only C. atricoxata is known; two larger females from Colombia have been examined but are not distinct other than size, but the poor condition of the specimens limits their evaluation. No males are known from Colombia, which makes species assessment with genitalia impossible. Diagnosis. A darkened, mottled appearance. Overall a squat species with robust legs and a short, broad pronotum and abdomen. The wings are marked with highly contrasting white or greenish and nearly black or dark brown markings. The lateral margins of the abdomen visible in females when the wings are folded over the abdomen.
Head (Fig. 46E): Juxta-ocular protuberances small; the vertex between the parietal sutures is straight; vertex lower than the dorsal margin of the eyes. Frontal suture with a faint curved carina. Ocelli small on protruding cuticular mounds. Head is dark brown with fine black speckling across the vertex and juxta-ocular protuberances, the frons, and parts of the clypeus.  Figure 36. Corticomantis atricoxata (Beier, 1931), dorsal habitus: A male from Costa Rica (USNM 026) B female from Costa Rica (USNM 003).
Pronotum (Fig. 50I): Short and broad with a less defined supra-coxal bulge; dorsal surface with tubercles of varying size and density. Broad prozone with lateral margins that taper anteriorly, the anterior margin rounded; a central depression medially on the dorsal surface. Metazone with lateral margins that are nearly parallel anteriorly, but taper dramatically to the narrowest point (the midpoint of the metazone) before becoming parallel anterior to the posterior terminus; the posterior margin rounded overall with the medial region being straight; posterior region with two blunt protrusions on each side of the medial line; the dorsal surface of the posterior half of the metazone depressed. Mostly pale in coloration with black speckling; two dark marks laterally on the posterior surface of the metazone.
Prothoracic Legs: Femur robust with a slightly concave dorsal margin distally, the proximal half slightly convex. Pale to dark banding on posterior (external) surface of femur, sometimes not well defined and degenerating into pale and dark speckling ventrally; anterior (internal) surface mostly black, but with pale regions dorsally; ventral surface mostly black with a pale margin just inside the posteroventral spines. Posterior prothoracic femoral genicular spine nearly the same length as the posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibia with pale and dark banding on the anterior and posterior surface, the ventral surface mostly pale. Prothoracic coxae black anteriorly (internal side) with a small pale marking in the proximal half along the dorsal margin.
Meso-and Metathoracic Legs: Femora with a pronounced dorsal and ventral (posterior) carina; posterior (upper) surface with two carinae, one running the nearly entire length, fading proximally and positioned ventrally to the dorsal carina, the other in the distal third and positioned parallel and between the previous and the ventral carina. Mesotarsi with first segment shorter than remaining segments combined. Metatarsi with first segment the same length as the remaining segments combined.
Wings: Forewings mottled with contrasting regions of brown, white, green, and black; the proximal quarter dark, then fading dramatically to a mottled white with a darkened spot on and around the pterostigma; the distal quarter of the wing mottled irregularly with half greenish white and half brownish black; the costal vein alternating from pale to dark its entire length while the costal region is mostly pale with banding in the distal quarter, but can be banded throughout its length; the costal region wide relative to the wing length, the width between 6.5-7.5% the total wing length. The forewings may be colored asymmetrically, one being mottled as described above while the other is blackened. Hindwings opaque and smoky; the terminus of the discoidal region projecting slightly beyond the distal margin of anal region, almost fitting within the outer margin of the wing.
Abdomen: Widening from first segment until the beginning of the distal half (segments 5-6) when the lateral margins narrow to the terminus, the middle being the broadest region. Tergites without pointed posterolateral projections. Supra-anal plate transverse with a blunt terminus. Subgenital plate of male with a rounded, slightly irregular terminus; without styli.
Genital Complex (Fig. 53E.1-E.3): Dextral genitalia with the main body of ventral left sclerite (L4A) with a prominent, heavily sclerotized distal process (pda) that forms a half circle, the terminus with a posterior orientation (Fig. 53, E.1 folded out and  (Piza, 1965), and Corticomantis atricoxata (Beier, 1931), dorsal habitus of types and labels. Fuga fluminensis (Piza, 1965): A holotype female (DZES) B labels. Corticomantis atricoxata (Beier, 1931): C holotype female from Costa Rica (ZMUH) D labels. 1999: 10; Salazar E. 2002: 124;Ehrmann 2002: 207;Otte and Spearman 2005: 133;Agudelo et al. 2007: 116, 142. Liturgusa mesapoda: Agudelo 2004  Taxonomic history. One of the earlier specimens described (Westwood 1889), H. mesopoda has always been included within Liturgusa. However, the examination of the holotype while treating the broader diversity of Liturgusa has revealed the species actually fits within the closely related genus Hagiomantis Saussure & Zehntner, 1894. Although the species is moved into Hagiomantis, the broader genus is not treated within this study, but H. mesopoda is redescribed herein. Two type specimens labeled as L. mesopoda from the Oxford University Museum of Natural History (OUMNH Type 452 2/3 and 3/3) were also examined and found to be L. cayennensis. The history of these two specimens is not known, but they should not be considered as types of H. mesopoda and may have been mislabeled at some point in the past. Further, Westwood (1889: 30) references only the holotype from St. Laurent de Maroni, French Guiana, and not any additional type material.
Diagnosis. Large species with a dark dorsal habitus, the coloration with highly contrasting light and dark mottling on the forewings. Numerous tubercles present on the posterior (external) surface of the prothoracic femora, the mesofemora, and the metafemora.
Head (Fig. 46G): Transverse with large, rounded eyes projecting outside the profile of the head both laterally and anteriorly (the anterior margin of the eyes anterior to the  central surface of the head). Juxta-ocular protuberances present, the lateral third being the most pronounced; the vertex between the parietal sutures is slightly concave; vertex lower than the dorsal margin of the eyes. Frontal suture with a faint curved carina, forming sharp angle medially. Two slightly protruding mounds are located symmetrically between the lateral sections of the frontal suture and the parietal sutures. Ocelli small with the central ocellus oriented anteriorly while the lateral are oriented anterolaterally at the 45 degree angle from the central axis of the head; protruding slightly on cuticular mounds. Frons narrowed between the antennal insertion sites and depressed below the central ocellus; a transverse carina present below the central ocellus, running from lateral margins under the antennal insertion sites medially in a dorsally oriented curve. Upper margin of clypeus convex, lower margin slightly concave; a central protruding ridge; the lateral margins tapering, widest at the upper margin. Labrum with minimal sculpting and a rounded terminus; lateral margins widening ventrally and concave. Antennae filiform and with rare setae, pale proximally and fading to black distally. Varying levels of black markings across the anterior surface of head. Palpi are pale.
Pronotum (Fig. 50K): Elongate with a less defined supra-coxal bulge; dorsal surface with numerous tubercles; prozone long with parallel lateral margins prior to a rounded anterior margin; metazone long with sweeping concave lateral margins that   exhibit small denticles; posterior margin of the metazone straight; the dorsal surface of the posterior half of the metazone depressed. Supra-coxal sulcus strongly defined. The lateral margins of the pronotum slightly expanded to form a small ledge. Colored with black and pale markings that vary across specimens.    (Beier, 1931): E male F female. H. mesopoda (Westwood, 1889): G male H female.   Prothoracic Legs: Femur robust with a slightly concave dorsal margin distally, the proximal third convex; anteroventral and posteroventral spines well developed; line of small, pointed tubercles running medially of the posteroventral spines; anteroventral spines black, posteroventral spines pale basally with black terminus. A strongly pronounced continuous carina running from distal terminus of femur along dorsal margin to the base, circling the posterior (external) surface of the proximal end and running  (Beier, 1931): I male J female. H. mesopoda (Westwood, 1889): K male L female. along the ventral margin at the base of the posteroventral spines. Pale to dark banding on posterior (external) surface of femur with numerous tubercles; anterior (inner) surface pale with femoral brush colored black; ventral surface pale. Well developed femoral pit on the ventral surface to accommodate terminal posteroventral tibial spine positioned on the lateral margin between the proximal two posteroventral spines, pushing the margin outward; pit is pale. Posterior prothoracic femoral genicular spine much smaller than posteroventral spines, originating distal to the beginning of the genicular lobe. Prothoracic tibial spines robust; the posteroventral spines with the first and second most proximal and fifth through seventh shorter than the slightly longer proximal third, fourth, and terminal spines; the anteroventral spines longest at distal end and shortening proximally. Prothoracic tibiae with a line of pronounced tubercles on the posterior surface, extending almost the entire length. Tarsi banded with pale and dark coloration. Prothoracic coxae smooth with varying black markings on the posterior and ventral surface, the anterior surface entirely pale.
Meso-and Metathoracic Legs: Long and slender with pale to dark banding on the femur and tibia. Femora with a pronounced ventral carina and a faint dorsal carina; posterior (upper) surface with pronounced tubercles. Tibiae banded with alternating Pronotum (Fig. 50L): Metazone with sharply rounded posterior corners caused by straightening of the lateral margins in the posterior terminus connecting to a straight posterior margin.
Prothoracic Legs: As described for males. Meso-and Metathoracic Legs: As described for males. Wings: As described for males. Abdomen: Elongate, gradually widening until the third to last tergite where it narrows dramatically, that last three segments being compressed. Tergites with small, pointed posterolateral projections. Supra-anal plate nearly square with a blunt, straight terminus.

Summary of taxonomic organization
When considering the phylogenetic results recovered by Svenson and Whiting (2009) the family-group, Liturgusidae, can not be considered a natural lineage. The name, stemming from the genus Liturgusa, could still apply to the Neotropical genera, but non-Neotropical genera would be taxonomically orphaned. The complexity of the polyphyly seen in Liturgusidae prevents simple taxonomic action since the higher-level relationships are still not robustly resolved to safely place non-Neotropical genera within established taxa. Therefore, these actions are avoided here as to not further destabilize the taxonomic organization. Listed here is the current taxonomic organization of the Neotropical Liturgusidae as well as a list of non-Neotropical genera that will eventually require placement within other family-groups. Interestingly, recent phylogenetic results stemming from new work on Neotropical mantises indicate that Gonatista Saussure, 1869 is not recovered with the Neotropical Liturgusidae, its fait currently unknown for higher-level placement (Rivera and Svenson, unpublished data).