ZooKeys 390: 1–214, doi: 10.3897/zookeys.390.6661
Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini)
Gavin J. Svenson 1,†
1 Department of Invertebrate Zoology, Cleveland Museum of Natural History, 1 Wade Oval Drive, Cleveland, Ohio 44106

Corresponding author: Gavin J. Svenson (gsvenson@cmnh.org)

Academic editor: L. Penev

received 22 November 2013 | accepted 29 January 2014 | Published 18 March 2014
(C) 2014 Gavin J. Svenson. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Svenson GJ (2014) Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini). ZooKeys 390: 1–214. doi: 10.3897/zookeys.390.6661

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.

Keywords

Mantodea, praying mantis, Liturgusa, Neotropical, taxonomy, new species

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, Liturgusa cayennensis, and created a new genus, Liturgousa, for his new species as well as Mantis annulipes. Saussure (1872: 260) later described Liturgusa 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).

In the same year, three new species were described, Liturgusa lichenalis, Liturgusa superba, and Liturgusa nubeculosa by Karl Eduard Adolph Gerstaecker (1889: 52–56) and Liturgusa mesopoda by John Obadiah Westwood (1889: 30). The publication by Westwood came slightly after that of Gerstaecker’s and included Mantis annulipes, Liturgusa cayennensis, Liturgusa lichenalis, Liturgusa nubeculosa, and his own new species Liturgusa mesopoda, which was the most complete reference to the species included within the group at the time.

The next species, Liturgusa maya, was described by Saussure and Zehntner (1894: 160) only as a variant of Liturgusa cayennensis, but was later elevated to a species by Samuel Hubbard Scudder (1901: 159).

William Forsell Kirby (1904: 271) included six species within Liturgusa: Liturgusa annulipes, Liturgusa lichenalis, Liturgusa cayennensis, Liturgusa maya, Liturgusa mesopoda, and Liturgusa malagassa. The last being a species from Madagascar and not related to the Neotropical Liturgusini (Svenson and Whiting 2009) and no longer included within the genus. Kirby also included Liturgusa nubeculosa, Liturgusa superba, and Liturgusa surinamensis within Hagiomantis without providing a justification for the move. Kirby furthermore fixed the type species for the genus to Liturgusa cayennensis Saussure, 1869.

Franz Werner (1906: 372) described Liturgousa orientalis from Tanzania, but later this species was moved to Dactylopteryx Karsch, 1892 with other species from the Afrotropical region.

Lucien Chopard (1911: 323, 1916: 164) recorded Liturgusa mesopoda, Liturgusa cayennensis, and Liturgusa annulipes from French Guiana, but his identification of Liturgusa annulipes is doubtful considering its currently recognized distribution.

Ermanno Giglio-Tos (1914: 77–78) described Liturgusa peruviana from Peru and Liturgusa parva from Brazil, citing that Liturgusa parva was similar to Liturgusa annulipes.

Morgan Hebard (1919b: 134) listed Liturgusa mesopoda as being from Jimenez Colombia, but the specimen Hebard examined is actually a female of a very large new species described herein (Liturgusa stiewei sp. n.), which may be why he identified it as the typically large Liturgusa mesopoda. Hebard (1922: 337) referenced a number of specimens of Liturgusa cayennensis from Central America, but this identification was probably incorrect (see below). Hebard (1924: 131) listed Liturgusa cayennensis as being from Ecuador, but the specimen was not located in the Academy of Natural Sciences of Drexel University, Philadelphia. In addition, the species location and identification records presented by Hebard (1929, 1932, 1933) are also not verifiable since the specimens could not be located.

Ermanno Giglio-Tos (1927: 292–295) treated the genus Liturgusa with a redescription of the genus, species redescriptions, and a species level key. Included within the work are eight species: Liturgusa peruviana, Liturgusa cayennensis, Liturgusa mesopoda, Liturgusa nubeculosa, Liturgusa annulipes, Liturgusa parva, and a new species, Liturgusa charpentieri. His new species was based on “non Mantis annulipes” specimens cited by Toussaint von Charpentier (1843). He also synonymized Liturgusa lichenalis with Liturgusa annulipes without justification or presentation of supporting observations. Giglio-Tos also cited Mantis annulipes as the type species for the genus. Interestingly, the species key provides a distinction between species with 7 or 8 posteroventral tibial spines as is done herein. However, the key is deficient in the characters used and includes Liturgusa nubeculosa within the 8 posteroventral spine group, which it does not exhibit.

Max Beier (1931: 14) described Liturgusa atricoxata based on a female specimen from the Zoological Museum in Hamburg (now named Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Germany). Then, Beier (1935: 11) provided a description of the genus as well as a list of nine species of Liturgusa that included Liturgusa peruviana, Liturgusa cayennensis, Liturgusa maya, Liturgusa charpentieri, Liturgusa atricoxata, Liturgusa mesopoda, Liturgusa nubeculosa, Liturgusa annulipes, and Liturgusa parva.

James Abram Garfield Rehn (1935) recognized early that the eight species included within Liturgusa were difficult to identify and that the group needed attention. Rehn went further to discredit the actions of Giglio-Tos in describing Liturgusa peruviana, which Rehn stated as an obvious junior synonym to Liturgusa nubeculosa. Rehn also examined the type of Liturgusa lichenalis to discredit Giglio-Tos’ action that synonymized the species with Liturgusa annulipes. Rehn dismissed the key provided by Giglio-Tos as it relies on color and misstated the characters of a number of species. Rehn also outlined the differences of Liturgusa maya and Liturgusa cayennensis, by which Liturgusa maya was described as a variant. He separated the two with morphological observations as well as stated that Liturgusa maya is mainly from Central America but can range into South America, while Liturgusa cayennensis is known from French Guiana. Rehn (1935: 202) also referenced specimens from Trinidad and identified them as Liturgusa maya, these turning out to be a new species described herein (Liturgusa trinidadensis sp. n.). Rehn (1935: 199–201) treated a species identified as Liturgusa annulipes, which later turned out to be Liturgusa cursor (see below). Rehn (1935: 204) also suggested that Liturgusa atricoxata may not actually be a member of the Liturgusa, but of another, possibly undescribed genus, a conclusion reached herein.

Marcello La Greca (1939: 2–5) thoroughly described the highly distinct species Liturgusa guyanensis with illustrations for two females collected in 1931 from Babooncamp (located in Guyana).

James Abram Garfield Rehn (1950) described two new species from Central America, one being previously treated as Liturgusa annulipes (Rehn 1935), but now named Liturgusa cursor (Rehn 1950: 369). The second species, Liturgusa actuosa (Rehn 1950: 377), was also described from Central America from specimens collected on Barro Colorado Island in Panama. Extremely thorough descriptions were provided along with figures, measurement data, and habits.

In addition to describing a new species of Liturgusa, Salvador de Toledo Piza Jr. described two new Hagiomantis species from Brazil including Hagiomantis fluminensis (1965) and Hagiomantis parva (1966). Years later he also described Liturgusa sinvalnetoi (1982: 94) from Piracicaba, Brazil. All three descriptions were brief, but did include measurement data as well as locality information.

Paulo S. Terra (1995) treated Liturgusa with a redescription as well as providing a list of 14 species in his study on Neotropical Mantodea. Included were Liturgusa actuosa, Liturgusa annulipes, Liturgusa atricoxata, Liturgusa cayennensis, Liturgusa charpentieri, Liturgusa cursor, Liturgusa guyanensis, Liturgusa lichenalis, Liturgusa maya, Liturgusa mesopoda, Liturgusa nubeculosa, Liturgusa parva, Liturgusa peruviana, and Liturgusa sinvalnetoi. Limited distributional information was also included.

Francisco J. Cerdá (1996: 75–76) listed three species in his treatment of Mantodea of Venezuela including Liturgusa nubeculosa, Liturgusa maya, and Liturgusa cayennensis. He also provided a description of the genus.

Julián A. Salazar E. (1998: 105, Fig. 5) listed Liturgusa charpentieri as a species found in Colombia, but based on the figure provided it resembles one of the new species described herein (Liturgusa krattorum sp. n. or Liturgusa algorei sp. n.). Salazar (1999: 10) also listed Liturgusa charpentieri and Liturgusa mesopoda from Para, Colombia, but the identification of Liturgusa mesopoda is doubtful based on that species being distributed within and around French Guiana.

In his dissertation, Lauro José Jantsch (1999: 47–48), listed both spellings of Liturgusa, but attributed the later spelling to Gerstaecker rather than Stål. Jantsch also provided a genus description and a list of 12 included species matching those presented by Terra (1995), but with Liturgusa lichenalis as a junior synonym of Liturgusa annulipes as well as citing Liturgusa peruviana (spelled as Liturgusa peruana) as a junior synonym of Liturgusa nubeculosa.

Jean-Michel Maes and Roger Roy (2000: 61) listed only one species from Nicaragua, Liturgusa maya.

Francesco Lombardo and Barbara Agabiti (2001: 90, 96–97) listed four species from Ecuador, three being new records for the country, which include Liturgusa cayennensis, Liturgusa charpentieri, Liturgusa maya, and Liturgusa peruviana.

Julián A. Salazar E. (2002: 124) listed three species from Para, Colombia including Liturgusa cayennensis, Liturgusa charpentieri, and Liturgusa mesopoda. This list reflects previous lists Salazar provided (1999) with the addition of Liturgusa cayennensis.

Reinhard Ehrmann (2002: 206–207) provided a genus description and a list of 14 described species of Liturgusa that includes Liturgusa actuosa, Liturgusa annulipes, Liturgusa atricoxata, Liturgusa cayennensis, Liturgusa charpentieri, Liturgusa cursor, Liturgusa guyanensis, Liturgusa lichenalis, Liturgusa maya, Liturgusa mesopoda, Liturgusa nubeculosa, Liturgusa parva, Liturgusa peruviana, and Liturgusa sinvalnetoi. Ehrmann also included an abbreviated bibliography for synonyms, type repository and sex, type locality and rough distribution for each species. As an aside, Ehrmann (2002: 163–164) listed Hagiomantis fluminensis Piza, 1965, Mantis ornata Stoll, 1813, Hagiomantis pallida Beier, 1942, Hagiomantis parva Piza, 1966, Liturgusa superba Gerstaecker, 1889, and Liturgusa surinamensis Saussure, 1872 for Hagiomantis.

Antonio Arnovis Agudelo Rondón (2004: 55) listed four species from Colombia, mostly reflecting previous lists by Salazar (1999), but with the addition of Liturgusa maya for the first time.

Daniel Otte and Lauren Spearman (2005: 132–133) included 14 species in their catalog identical to those listed by Ehrmann (2002), but provided additional bibliographic references for the genus and each species, type repository, and limited distributional information.

Antonio Arnovis Agudelo Rondón, Francesco Lombardo, and Lauro José Jantsch (2007: 116, 141–142) listed 13 valid species in their checklist of Neotropical Mantodea. Liturgusa peruviana was synonymized with Liturgusa nubeculosa.

David Yager and Gavin Svenson (2008: 556) included a Liturgusa species in a molecular based phylogeny for the first time. This was soon followed by Gavin Svenson and Michael Whiting (2009) including Liturgusa maya and Liturgusa tessae sp. n. in their molecular study. The two species were placed as sister to Hagiomantis within a large diversity of Neotropical Mantodea.

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 Liturgusa 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.

Figure 1.

Liturgusa habitus of live specimens: A Liturgusa cursor Rehn, 1935, female from Nicaragua B Liturgusa kirtlandi sp. n., female from Bolivia C Liturgusa nubeculosa Gerstaecker, 1889, female from Bolivia D ootheca of unknown species of Liturgusa.

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 Liturgusa algorei sp. n. in Peru). Larger females of Liturgusa 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 the forest. Specimens have been observed in higher regions of tree (Liturgusa nubeculosa, Liturgusa fossetti sp. n., and Liturgusa krattorum sp. n.) as well as within the crown of trees (Liturgusa 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 (Liturgusa 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 (Liturgusa nubeculosa and Liturgusa maya) and Nicaragua (Liturgusa fossetti sp. n. and Liturgusa 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, Liturgusa maya was readily collected in seasonal deciduous forests in western Nicaragua during the dry season, which is interesting considering that Liturgusa 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 Liturgusa 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 Liturgusa maya with a specimen collected in Mexico, a little north of Mexico City near the eastern coast. The southernmost record of Liturgusa are those of Liturgusa nubeculosa and Liturgusa kirtlandi sp. n. from just west of Santa Cruz, Bolivia. Species included within the newly created genera, Fuga gen. n. and Velox gen. n., are entirely restricted to southeast Brazil. This range is disjunct from the most eastern ranging species of Liturgusa.

A few of the species have broad ranges, covering thousands of square kilometers across the Amazon Basin (Liturgusa nubeculosa and Liturgusa tessae sp. n.) or with an extended north to south range from Mexico to southern Peru (Liturgusa maya). Most species have been found to be relatively restricted in their distribution, some only occurring within one country, island or even region of a country. Regional restriction is the consistent pattern across the group and is congruent with what would be expected of species that do not appear to fly or disperse readily. However, the broad ranges of the three aforementioned species warrants further investigation. A population genetics study would likely lead to even more diversity through discovery of cryptic species.

Chromosomes

The chromosome composition of three species of Liturgusa has been characterized in multiple works by Sally Hughes-Schrader and her students (Hughes-Schrader 1943, 1948, 1950, 1951, 1953). Her interest in Mantodea for this work was likely due to a collaboration with James A.G. Rehn of the Academy of Natural Sciences of Drexel University, Philadelphia. Liturgusa actuosa was shown to have 23 chromosomes, Liturgusa cursor to have 33, and Liturgusa maya to have 17. All three species have a XO sex chromosome composition for males.

Revision

This morphologically based revision assembled a large representation of Liturgusa specimens in order to address the impossibility of identifying most of the described species. With few keys present in the literature, most specimens of Liturgusa were either undetermined or misidentified. This lack of information for the genus has hampered our knowledge about their true diversity. In addition, based on their behavior, habitat usage and abundance in many environments they would make ideal study subjects for many types of ecological studies. In fact, a similar but unrelated Australasian bark mantis genus, Ciulfina, has received such attention with both taxonomic and ecological research, both of which have produced highly interesting results (Attard et al. 2009; Holwell 2008; Holwell and Herberstein 2010a, 2010b; Holwell et al. 2007, 2010; O’Hanlon 2011; Umbers et al. 2011).

The size of this study was not anticipated, but soon after gathering specimens and type material, it became clear that not only did synonymies exist, but a large number of new species were in need of description. The scope of this study then focused on the taxonomy of the genus alone with future plans to produce a robust morphological and molecular based phylogeny for the group.

The present study treats all described species of Liturgusa (11 species), describes 19 new species, identifies four synonymies, moves one species from Liturgusa to Hagiomantis, and creates three new genera (Fuga gen. n., Velox gen. n., and Corticomantis gen. n.) for species previously included in Liturgusa as well as Hagiomantis.

Methods

Specimens examined. A total of 522 specimens of Neotropical Liturgusini were examined from 25 collections and museums. Approximately half were collected previous to 1950, thus incorporating a significant historical sampling from within the group. Each specimen was assigned a unique code to track locality, measurement data, and images. These codes will remain on the specimen pins, but it can’t be guaranteed that specimens will permanently retain the assigned code after return of loaned material. All data for examined specimens appear in tables prior to the descriptions, which include sex, type status, country of origin, locality data, georeferenced coordinates, and project code that includes repository abbreviations. In addition to the project codes, new database codes have been added to specimens from the National Museum of Natural History, Smithsonian Institution. A spreadsheet version of material examined tables can be downloaded at http://mantodearesearch.com.

Georeferencing. A specimen level map was created within Google Earth, each waypoint with a species specific marker and a description that includes specimen codes, label data, and type status. The KML file can be downloaded at http://mantodearesearch.com and opened in Google Earth and will enable users interactive capabilities when displaying distribution data. Specimens without GPS data included on the label were georeferenced using online map databases and Google Earth. All locality information presented in the material examined tables was reformatted and input into templates for conversion to KML format by Earth Point © (http://www.earthpoint.us/ExcelToKml.aspx).

Descriptive conventions and character systems. The species treated within this study are all extremely similar based on external morphology. However, key external features exist that easily separate the Neotropical genera. Species treatments within this study provide a brief diagnosis, new natural history observations, and verbal character descriptions stemming from the anterior surface of the head, the dorsal surface of the pronotum, the prothoracic leg (spine terminology following Wieland 2008, 2013), the wings, and the abdomen. The verbal descriptions are provided for males and females as well as the ootheca for one species. When females match male features, they are excluded from the female description. Therefore, only characters that differ slightly from the males are presented in the female descriptions, all other characters can be assumed to match the male.

Male genital complex. To extract the genital complex, terminal abdominal segments were dissected and placed in a hot weak KOH solution for 30 min to dissolve muscle tissue. Cleared genital structures were disarticulated and slide mounted with 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.

  5. Prozone length = anterior margin of pronotum to center of supra-coxal sulcus.

  6. 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 (50 mm, 100 mm, and MP-E 65 mm), 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.

Figure 2.

Liturgusa fossetti sp. n., dorsal habitus illustration: A holotype male from Panama (CLEV GSMC003836) B allotype female from Nicaragua (CLEV GSMC003425).

Specimen deposition. Examined specimens are deposited or reside in the following institutions. New type material was deposited in accordance to specimen ownership with slide mounted male genital complex included. Museum codes are used throughout for brevity.

AMNH American Museum of Natural History, New York, NY, USA

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA

BMNH The Natural History Museum (British Museum Natural History), London, UK

CAS California Academy of Sciences, San Francisco, CA, USA

CJASE Collection Julian A. Salazar E, Manizales, Colombia

CLEV Cleveland Museum of Natural History, Cleveland, OH, USA (GSMC codes are all CLEV database record codes for the collection of Mantodea)

DZES Universidade de Sao Paulo, Piracicaba, Brazil

EMAU Ernst-Moritz-Arndt-Universität Greifswald, Germany

FMNH Field Museum of Natural History, Chicago, IL, USA

GSMC Gavin Svenson Mantodea Collection of Research, Cleveland Museum of Natural History, Cleveland, OH, USA

JAC Collection of J. Amith

LEM Lyman Entomological Museum, McGill University, Quebec, Canada

MALUZ Museo de Artrópodos de la Universidad del Zulia, Maracaibo, Venezuela

MEKRB Museo de Entomología Klaus Raven Buller, Universidad Nacional Agraria La Molina, Lima, Peru

MHNG Muséum d’Histoire naturelle, Geneva, Switzerland

MLBM Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah, United States of America

MNHN Muséum national d’Histoire naturelle, Paris, France

MNKM Museo de Historia Natural Noel Kempff Mercado, Santa Cruz, Bolivia

MSMC Martin Stiewe Mantodea Collection, London, United Kingdom

NYSM New York State Museum, Albany, NY

OUMNH Oxford University Museum of Natural History, Oxford, United Kingdom

ROM Royal Ontario Museum, Toronto, Canada

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany

USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany

ZMUH Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Germany

Taxonomic treatment
Key to genus
1 Prominent tubercles distributed across the posterior (upper) surface of the meso- and metafemora Hagiomantis Audinet Serville, 1838
Posterior (upper) surface of the meso- and metafemora smooth 2
2 Number of posteroventral tibial spines 7, in extremely rare cases females may have 8 on one or both tibiae. The second most proximal posteroventral spines larger than others, excluding the most distal spine Liturgusa Saussure, 1869
Number of posteroventral tibial spines 8. The third most proximal posteroventral spine larger than others, excluding the most distal spine 3
3 The prothoracic femoral pit that accommodates the terminal posteroventral tibial spine positioned between the two most proximal posteroventral spines, extending laterally abutting the posterior margin of the femur. Subgenital plate of male with pronounced styli Velox gen. n.
The prothoracic femoral pit that accommodates the terminal posteroventral tibial spine positioned between the first most proximal posteroventral spines and the most distal discoidal spine. Subgenital plate of male without styli 4
4 Tubercles distributed across dorsal surface of the pronotum. Pronotum shape measure (pronotum width at the supra-coxal bulge over total length of pronotum) of male and female 0.51 or greater (observed range 0.51–0.55); pronotum short and broad. Distributed entirely in Central America or at most in northern South America (two specimens are from Colombia) Corticomantis gen. n.
Dorsal surface of the pronotum is smooth or at most with very small tubercles widely dispersed. Pronotum shape measure (pronotum width at the supra-coxal bulge over total length of pronotum) of male and female 0.49 or less (observed range 0.33–0.49); pronotum more elongate. Distributed in eastern Brazil and not extending into the Amazon basin Fuga gen. n.

Clave Para Género (translation by Julio Rivera)

1 Meso- y metafemures con tuberculos prominentes distribuidos a lo largo de su superficie posterios (dorsal) Hagiomantis Audinet Serville, 1838
Meso- y metafemures con superficie posterios (dorsal) lisa 2
2 Tibias anteriores con 7 espinas posteroventrales, aunque excepcionalmente las hembras pueden mostrar 8 espinas en una o ambas tibias. La 2da y 3ra espinas tibiales posteroventrales proximales más largas que las demás, excepto la más distal Liturgusa Saussure, 1869
Tibias anteriores con 8 espinas posteroventrales. La 3ra espina tibial posteroventral más proximal es más larga que las demás, excepto la más distal 3
3 La espina posteroventral más distal de la tibia anterior se inserta, cuando flexionada contra el fémur, en una depresión ubicada entre las dos espinas posteroventrales más proximales de este último; dicha depresión se extiende lateralmente hasta alcanzar el margen posterior del fémur. Placa subgenital del macho con styli prominente Velox gen. n.
La espina posteroventral más distal de la tibia anterior se inserta, cuando flexionada contra el fémur, en una depresión ubicada entre la primera espina posteroventral más proximal de este último y la espina discoidal más distal. Placa subgenital del macho sin styli 4
4 Superficie del pronotum con tubérculos. Medida de la forma del pronotum (i.e. ancho del pronotum a la altura de la dilatación supracoxal dividido por la longitud total del pronotum) del macho y hembra igual o superior a 0.51 (rango observado 0.51–0.55); pronotum corto y ancho. Distribución estrictamente Centroamericana o apenas hacia el norte de Sudamérica (dos especímens estudiados fueron obtenidos en Colombia) Corticomantis gen. n.
Superficie del pronotum lisa o a lo mucho con túberculos muy pequeños y ampliamente dispersos. Medida de la forma del pronotum (i.e. ancho del pronotum a la altura de la dilatación supracoxal dividido por la longitud total del pronotum) del macho y hembra igual o menor a 0.49 (rango observado 0.33–0.49); pronotum más bien alargado. Distribución en el este de Brazil, no ocurren en la Cuenca del Amazonas Fuga gen. n.
Liturgusa Saussure, 1869

http://species-id.net/wiki/Liturgusa

Liturgousa: Saussure 1869: 55, 62; Brauer 1870: 92; Saussure 1870: 225; Brauer 1871: 189; Saussure 1871a: 29, 51, 205; Saussure 1871b: 100; Saussure 1872a: 20, 53, 156; Saussure 1872b: 259; Stål 1877: 50; Westwood 1889: 4, 30; Brunner de Wattenwyl 1893: 63, 225; Saussure and Zehntner 1894: 129, 159; Saussure and Zehntner 1895: vii, 155, 157; Badenoch 1899: 39; Scudder 1901: 159, 407, 419; Waterhouse 1902: 202; Rehn 1903: 6; Kirby 1904: 271; Bruner 1906: 143; Werner 1906: 372; Werner 1908: 39; Werner 1909: 77–78; Chopard 1911: 323; Carl 1914: 148; Chopard 1916: 164; Werner 1916: 257, 274; Caudell 1918: 5; Hebard 1919a: 31; Hebard 1919b: 134; Hebard 1924: 131; Giglio-Tos 1927: 292 (“err. transscript.”), 699; Hebard 1929: 399; Hebard 1932: 211; Hebard 1933: 29; Rehn 1935: 172, 198–199, 201, 203–204, pl. 8, Figs 4–5; Hesse 1937: 108, 578; Tinkham 1937: 490–491; Neave 1939: 978; Hughes-Schrader 1943: 266, 280, 282–283, 290, 294, 296–297, Table 1, Figs 19–28; Hughes-Schrader 1948: 267; Rehn 1950: 369, 377; Hughes-Schrader 1950: 10–11, 13–14, 27, 38–39, 44–45, Table 1, Figs 9–11; Hughes-Schrader 1951: 178–181, 183–184, 186–187, Table 1–2, Figs 1–3; Beebe et al. 1952: 245–247, Fig. 2; Crane 1952: 259, 264, Figs 2, 3; Hughes-Schrader 1953: 544–554; Rehn 1954: 177, 179; Schrader and Hughes-Schrader 1956: 493–494, 496; Callan and Jacobs 1957: 201; Krombein 1963: 2; Henderson 1965: 206, 215; White 1965: 542; Marshall 1975: 318; Otte 1978: 76; Terra 1995: 53–54 (error); Cerdá 1996: 75–76, Fig. 1; Jantsch 1999: 47 (error transcript); Ehrmann 2002: 206 (Liturgousa err. descr.); Otte and Spearman 2005: 132, 481 (syn); Agudelo et al. 2007: 141 (syn); Roy and Cuche 2008: 8, 14, 21.
Liturgusa: Stål 1877: 3, 40; Gerstaecker 1889: 52–54 (idem); Bertkau 1889: 87; Giglio-Tos 1914: 77; Giglio-Tos 1916: 154; Giglio-Tos 1917: 64, 83; Carus 1921: 317; Hebard 1922: 337; Apolinar M. 1924: 47; Giglio-Tos 1927: 292; Beier 1931: 14; Brues and Melander 1932: 89, 644; Beier 1935: 6, 11; Rehn 1935: 198, 201, 204; Apolinar M. 1937: 226; La Greca 1939: 2, Fig. 1; Neave 1939: 978; Brues et al. 1954: 89, 878; Beier 1964: 943; Weidner 1964: 143; Beier 1968: 8, 14, 32; Marshall 1975: 322; Bazyluk 1977: 133, 169; Passerin d‘Entrèves 1981: 61; Piza 1982: 94; Terra 1995: 53–54, Figs 85–87; Salazar E. 1998: 105, Fig. 4; Edmunds and Brunner 1999: 282; Jantsch 1999: 19, 24, 30–31, 33, 35, 39, 47, Tables 4–6; Roy 1999: 30; Salazar E. 1999: 10; Maes and Roy 2000: 61; Salazar E. 2000: 67; Lombardo and Agabiti 2001: 90, 96–97; Salazar E. 2002: 121, 124; Ehrmann 2002: 26, 33, 206, 375; Agudelo and Chica 2002: 7, 20, 30, 36, 62, Fig. 8b; Agudelo and Chica 2003: 127, 130, 131, 132, 133, Tables 1, 3, Figs 1, 3, 7; Agudelo 2004: 44, 55, Table 3.1; Salazar E. 2004: 211, 213; Agudelo 2005: 3; Otte and Spearman 2005: 132, 481; Agudelo et al. 2007: 109, 116, 141; Medellín et al. 2007: 151; Roy and Cuche 2008: 8, 14, 21 (emendation by Stål); Yager and Svenson 2008: 556, 565; Wieland 2008: 158; Svenson and Whiting 2009: 503, Appendix S1; Wieland 2013: 22, 57, 87, 89, 130, 154, 158, 176, Figs 2, 4A, 20–21; Svenson in press.
Hagiomantis (partim): Kirby 1904: 271.
Liturguda: Jantsch 1991: 125.
Type species.

Liturgusa cayennensis Saussure, 1869 (Designation by Kirby 1904: 271).

Taxonomic history.

Spelling of genus. Two spellings of the genus name are present in the literature. The first is Liturgousa, established by Henri de Saussure in 1869 to include two species, Mantis annulipes Audinet Serville, 1838 and his newly described Liturgousa cayennensis Saussure, 1869. The second is Liturgusa, introduced by Carl Stål in 1877 while attributing the name to the original author, Saussure (1869). The name is derived from the Greek Liturgus (feminine form Liturga), meaning “celebrator of liturgy”, which indicates that Saussure’s original spelling of Liturgousa may have been a mistake, but under Article 32.5.1 of the ICZN (International Code of Zoological Nomenclature 4th edition) “incorrect transliteration or latinization, or use of an inappropriate connecting vowel, are not to be considered inadvertent errors” and thus is not demonstrably incorrect under Article 32.5 and stands as the correct original spelling under Article 32.2. The subsequent spelling of Liturgusa proposed by Stål appears to be an emendation, but since this emendation is applied to a correct original spelling and he also did not include a justification for his subsequent spelling (Art. 33.2.1), it is considered as an unjustified emendation under Article 33.2.3 of the Code. Under Article 33.2.3.1 the unjustified emendation (Liturgusa) becomes justified when it is in prevailing usage and is attributed to the original author and date. A comprehensive nomenclatural note pertaining to the spelling issue of Liturgusa has established such prevailing usage (Svenson in press), thus the unjustified emendation was determined as justified and is an available name that is used herein.

Type designation.

Upon creation of the genus Liturgusa by Saussure in 1869, two species, Mantis annulipes Audinet Serville, 1838 and Liturgusa 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 Liturgusa 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 Liturgusa cayennensis Saussure, 1869 as the type for the genus has not been uniform across taxonomic works (e.g. Ehrmann 2002 recognizes Mantis annulipes and Otte and Spearman 2005 recognizes Liturgusa 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.

Measurement Ranges: Male. Body length 18.59–29.16; forewing length 10.74–17.91; hindwing length 8.12–14.19; pronotum length 5.03–8.84; prozone length 1.55–2.49; pronotum width 1.90–3.18; pronotum narrow width 1.27–2.47; head width 4.36–6.34; head vertex to clypeus 1.55–2.67; frons width 1.39–2.39; frons height 0.53–0.93; prothoracic femur length 5.40–8.10; mesothoracic femur length 3.32–11.15; mesothoracic tibia length 4.78–8.83; mesothoracic tarsus length 4.33–8.07; metathoracic femur length 6.61–11.64; metathoracic tibia length 6.05–12.00; metathoracic tarsus length 6.24–11.52; pronotal elongation measure 0.26–0.34; pronotal shape measure 0.28–0.49; head shape measure 0.36–0.44; frons shape measure 0.30–0.47. Female. Body length 14.64–52.03; forewing length 12.36–26.96; hindwing length 9.29–21.20; pronotum length 5.16–13.12; prozone length 1.57–3.69; pronotum width 2.19–4.49; pronotum narrow width 1.43–3.09; head width 4.42–7.91; head vertex to clypeus 1.78–3.38; frons width 1.51–3.23; frons height 0.52–1.23; prothoracic femur length 5.22–12.24; mesothoracic femur length 6.47–14.62; mesothoracic tibia length 4.83–12.35; mesothoracic tarsus length 4.56–10.74; metathoracic femur length 6.56–14.87; metathoracic tibia length 6.90–17.08; metathoracic tarsus length 6.38–14.80; pronotal elongation measure 0.23–0.33; pronotal shape measure 0.29–0.52; head shape measure 0.38–0.47; frons shape measure 0.30–0.44.

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, 21A–B). 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.

Key to species
1 Pronotum shape measure (measure = pronotum width at the supra-coxal bulge / total length of pronotum) of male 0.435 or greater, of female 0.45 or greater; pronotum short and broad (Fig. 47A–G). Metazone shape metric (metric = pronotum width at the supra-coxal bulge / length of metazone) of male 0.63 or greater, of female 0.63 or greater; short, broad metazone 2, Cayennensis Group
Pronotum shape measure of male 0.43 or less, of female 0.43 or less; pronotum slightly to highly elongate (Figs 47H–T, 48, 49). Metazone shape metric of male 0.61 or less, of female 0.615 or greater 5
Cayennensis Group
2 Pronotum widened at the posterior end, the lateral margins protruding; distinct narrowing in the medial region of the metazone (Fig. 47E–F) 3
Pronotum with gradually narrowing lateral margins from supra-coxal bulge to posterior margin with no or very little central narrowing; posterior lateral margins not protruding and contained within the profile of the posterior terminus (Fig. 47A–D). Terminus of apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) broad at base, quickly tapering to sharp or blunt point; often triangular (Fig. 51A–B) 4
3 Lateral margins of male and female prozone parallel or slightly divergent anteriorly (Fig. 47E–F). Discoidal region of hindwing of male and female with orange or rust coloration fading to smoke distally and into the anal region (Fig. 6A–B). Terminus of apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) narrow and shaped like a finger (Fig. 51C.1). Distributed in northern Guyana and northern Brazil Liturgusa guyanensis La Greca, 1939
Lateral margins of the female prozone converging anteriorly (Fig. 47G). Discoidal region of hindwing of female (only females known) yellow on proximal 90% and fading to black distally; yellow extending into the anterior portion of the anal region with strongly defined transition from yellow to hyaline with a slight smoke color along the distal margin (Fig. 7A). Only known from southern Venezuela near the Sierra de la Neblina Liturgusa neblina sp. n.
4 Mesotarsi with first segment as long as the remaining segments combined. Distributed in eastern South America including French Guiana, Guyana, Suriname, and eastern Brazil. Lateral margins of female abdomen with slight or no tergal expansions Liturgusa cayennensis Saussure, 1869
Mesotarsi with first segment shorter than the remaining segments combined. Distributed in western South America including Colombia, Ecuador, Peru, and southern Venezuela. Lateral margins of female abdomen with distinct tergal expansions that produce a posterior oriented point Liturgusa lichenalis Gerstaecker, 1889
5 Pronotum shape measure of male 0.360 or greater, of female 0.370 or greater 6, Maya Group
Pronotum shape measure of male 0.355 or less, of female 0.36 or less; pronotum elongate and slender 15, Cursor Group
Maya Group
6 Forewing length of female 26–27 mm, of male 19–20 mm. Length of pronotum of female 11–12 mm, of male 9–10 mm. Known only from Colombia Liturgusa stiewei sp. n.
Forewing length of female 13–22 mm, of male 12–18 mm. Length of pronotum of female 6–10 mm, of male 5–8 mm 7
7 Pronotum highly sculpted in male and female with all of the following characteristics: prozone with highly convex margins; a defined supra-coxal bulge; medial region of metazonal lateral margins parallel or highly convex and bulging outward; posterior margins rounded in males, females with straightened posterior corners oriented 45 degrees from the central line of the body (Fig. 47E–F). Distributed only in Central America with records from Honduras, Guatemala, Belize, and Panama Liturgusa zoae sp. n.
Pronotum of male and female not highly sculpted with all of the following characteristics: prozone with lateral margins that are near parallel or tapering anteriorly; a moderately defined supra-coxal bulge; medial region of metazonal lateral margins concave or bulging slightly; posterior margins in males and females rounded with a central emargination (Liturgusa manausensis male with straightened posterior corners oriented 45 degrees from the central line of the body) (Figs 47H–T, 48C–D) 8
8 Relatively small with a size metric range (size metric = length of forewing × length of pronotum) of 63–68 for males and 88–91 for females 9
Relatively large with a size metric range of 73–132 for males and 97–187 for females 10
9 Prothoracic femora of female with posteroventral genicular spine positioned proximal to the beginning of the genicular lobe. Male size metric approximately 16–17 (size metric = [length of pronotum / width of pronotum at narrow] × width of head). Distributed in northern Peru with one record just across the border in Brazil. Liturgusa bororum sp. n.
Prothoracic femora of female with posteroventral genicular spine positioned distal to the beginning of the genicular lobe. Male size metric approximately 14. Distributed in northern Venezuela Liturgusa cura sp. n.
10 Hindwings of male and female with yellow, orange or rust coloration on at least the proximal third of the discoidal region, fading to black distally (Figs 10A–B, 16A–B) 11
Discoidal region of the hindwings of male and female black or darkly opaque with at most the anterior costal margin with a lighter pale or rust coloration (Figs 11A–B, 12A, 13A–B, 17A–B) 12
11 Hindwings of male and female orange or yellow colored on the proximal ¾ of the discoidal region, sharply transitioning to black distally; the anal region mostly black or darkly opaque with the proximal third matching the orange color of the discoidal region (Fig. 10A–B). Size metric (metric = length of prothoracic femora / length of forewing) for males 0.45 or greater, for females 0.47 or greater. Ventral left sclerite (L4A) without distal process (pda), smoothly rounded (Fig. 51F.1). Distributed in Central America with records from Panama, Costa Rica, and Nicaragua Liturgusa fossetti sp. n.
Hindwings of male and female orange or rust colored on the proximal ⅓ to ½ of the discoidal region, gradually fading to black distally; the anal region uniformly opaque black or smoky (Fig. 16A–B). Size metric for males 0.44 or less, for females 0.45 or less. Ventral left sclerite (L4A) with distal process (pda) (Fig. 51J.1–J.6) Liturgusa tessae sp. n.
Male genital dissection required for identification of Liturgusa manausensis, Liturgusa kirtlandi, Liturgusa trinidadensis, and Liturgusa maya.
12 Male pronotal shape metric 0.59 (metric = [length of prozone / length of pronotum] × pronotum width at the supra-coxal bulge). Apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) with a long, sharp point; ventral plate (pia) and ventral process (pva) of right phallomere (R1) large (Fig. 51H.1). Small species with asymmetrical forewing coloration, the contrast between dark and light highly distinct. Only males known and recorded only from Manaus, Brazil Liturgusa manausensis sp. n.
Pronotal shape metric for males 0.63 or greater, for females 0.77 or greater. Apofisis falloid (afa) of the main body of dorsal left sclerite (L4B) with a short, but sharp point; ventral plate (pia) and ventral process (pva) of right phallomere (R1) small (Figs 51G.1–G.4, 51I.1–I4, 52A.1–A.5) 13
13 Apical process (paa) of dorsal left sclerite (L4B) flat, forming an angled pedestal (Fig. 51G.1–G.4). Relatively large with a pronotum length range of 6.95–7.35 mm (7.18 average) for males and 8.16–8.70 mm (8.43 average) for females. Recorded only from central Bolivia Liturgusa kirtlandi sp. n.
Apical process (paa) of dorsal left sclerite (L4B) uniformly rounded (Figs 51I.1–I4, 52A.1–A.5). Relatively smaller overall with a pronotum range of 5.61–7.39 mm (6.25 average) for males and 6.55–8.99 mm (7.42 average) for females (only one male and two females of Liturgusa maya fall within the pronotum size range of Liturgusa kirtlandi) 14
14 Numerous tubercles in posterolateral corners of metazone in males and females. Antennae pale or gradually fade of a brown color in both sexes. Distal process (pda) of ventral left sclerite (L4A) with a sclerotized fold or slight projection centrally (Fig. 52A.1–A.5). Known only from Trinidad Liturgusa trinidadensis sp. n.
No tubercles present on pronotum of males and females. Antennae of both sexes fade rapidly to black just distal of the base. Distal process (pda) of ventral left sclerite (L4A) with a strongly defined central process that is angled approximately 30 degrees from the central axis of the hypophallus, the tip rounded (Fig. 51I.1–I4). Broadly distributed from Mexico to southern Peru on the western side of South America Liturgusa maya Saussure & Zehntner, 1894
15 Length of forewing of male less than 13.2 mm. Size metric (metric = length of forewing × length of pronotum) of male 92 or less, of female 125 or less. Ventral left sclerite (L4A) with a smooth, rounded terminus, lacking distal process (pda) (Fig. 52C.1–E.1) 16, Cursor Group A
Length of forewing of male 13.2 mm or greater. Size metric (metric = length of forewing × length of pronotum) of male 95 or greater, of female 130 or greater. Ventral left sclerite (L4A) with a distal process (pda) or serrated ridge (Fig. 52F.1–K.1) 18, Cursor Group B
Cursor Group A
16 Supra-anal plate of male and female highly transverse, about three times as wide as long. Forewings lacking strongly pronounce white spot near the proximal end (Fig. 19A–B). Apofisis falloid (afa) of the dorsal left sclerite (L4B) finger-like with the terminus shaped like a slightly broadened bulb (Fig. 52C.1–C.3). Distributed only in Central America with records in Nicaragua, Costa Rica, and Panama Liturgusa cursor Rehn, 1950
Supra-anal plate of male and female about twice as wide as long, the terminus rounded. Forewings with strongly pronounce white spot in the first proximal fifth, overlaying the bases of the media and cubitus veins (Figs 20A–B, 22A–B). Apofisis falloid (afa) of the dorsal left sclerite (L4B) triangular, terminating with a sharp point (Fig. 52D.1–E.1) 17
17 Anterior surface of forecoxae with a broad, black band medially in the proximal half as well as a black spot medially towards the distal terminus. Posterior margin of the prothoracic femora with a small spine between the most distal posteroventral spine and the genicular spine. Genicular spine positioned proximal to the beginning of the genicular lobe. Distributed in northern South America with records from French Guiana, but range could extend into Suriname and northern Brazil Liturgusa milleri sp. n.
Anterior surface of forecoxa pale, without black markings. Posterior margin of the prothoracic femora without defined spine between the most distal posteroventral spine and the genicular spine. Genicular spine positioned distal to the beginning of the genicular lobe. Endemic to the island of Dominica Liturgusa dominica sp. n.
Cursor Group B
18 Discoidal area of the hindwings red, rust, or orange color proximally and fading to black distally (Figs 26A–B, 29A). Forewings of male and female often with asymmetrical coloration; when present, one is red or rusty 19
Discoidal area of the hindwings uniformly dark or smoky colored (Figs 7B, 23A–B, 24A–B, 25B, 27A–B). Forewings of male and female with or without asymmetrical coloration; when present, one is darkened black or smoky 20
19 Posteroventral prothoracic femoral spines of males and females long, most proximal spine around 0.50 mm and 1 mm, respectively (length of prothoracic femora ~14 and ~8.5 times the length of the most proximal posteroventral prothoracic femoral spine, respectively). Apofisis falloid (afa) of the dorsal left sclerite (L4B) a slender point, appearing like a needle; terminus of ventral left sclerite (L4A) with a sharp, slightly curved distal process (pda), but it can be reduced to a shorter projection with a pointed tip (Fig. 52I.1) Liturgusa krattorum sp. n.
Posteroventral prothoracic femoral spines of male and female short, most proximal spine around 0.35 mm and 0.50 mm, respectively (length of prothoracic femora 18 and 15.5 times the length of the most proximal posteroventral prothoracic femoral spine, respectively). Apofisis falloid (afa) of the dorsal left sclerite (L4B) a broad triangle ending with a sharp point; terminus of ventral left sclerite (L4A) with a small blunt distal process (pda) (Fig. 52K.1) Liturgusa purus sp. n.
20 Length of pronotum of male less than 7.1 mm, of female less than 8 mm Liturgusa actuosa Rehn, 1950
Length of pronotum of male greater than 7.1 mm, of female greater than 8.5 mm 21
21 The length of the forewing is at most 1.80 times longer than the total length of the pronotum (observed range 1.67–1.80). Hindwings of male and female with a slight emargination between the discoidal and anal regions (Figs 7A, 25B) 22
The length of the forewing is at least 1.83 times longer than the total length of the pronotum (observed range 1.83–2.2). Hindwings of male and female with a prominent emargination between the discoidal and anal regions (Figs 25A–B, 27A–B) 23
22 The second most proximal posteroventral foretibial spine of female approximately 1.4 times the length of the third spine. Abdomen of female with slight posteriorolateral tergal projections that form a small posterior corner lip at most. Distributed in northern Venezuela with one record from northern Guyana Liturgusa cameroni sp. n.
The second and third most proximal posteroventral foretibial spines of females roughly the same length, the second being slightly longer in most specimens (not more than 1.2 times the length of the third spine). Abdomen of female with pronounced posterolateral tergal projections that form small tooth-like projections between 0.15 mm and 0.21 mm in length. Distributed in northern French Guiana Liturgusa maroni sp. n.
23 Mesotarsi of male with first segment shorter than the remaining segments combined. First tarsomere of female hindleg 3.5–8 times longer than the dorsal metatibial spur (spine relatively long). Distal process (pda) of ventral left sclerite (L4A) pointed, one side being serrated and heavily sclerotized; apofisis falloid (afa) of the dorsal left sclerite (L4B) with a broad, triangular terminus (Fig. 52J.1–J.2) Liturgusa nubeculosa Gerstaecker, 1889
Mesotarsi of male with first segment equal to remaining segments combined. First tarsomere of female hindleg 10–12 times longer than the dorsal metatibial spur (spine relatively short). Distal process (pda) of ventral left sclerite (L4A) pointed, the tip rounded; apofisis falloid (afa) of the dorsal left sclerite (L4B) with slender, pointed terminus (Fig. 52G.1–G.2) Liturgusa algorei sp. n.
Clave Para las Especies (translation by Julio Rivera)
1 Medida de la forma del pronotum (medida = ancho del pronotum a la altura de la dilatación supracoxal dividido por la longitud total del pronotum) del macho igual o mayor a 0.435, el de la hembra igual o mayor a 0.45; pronotum corto y ancho. Medida de la forma de la metazona (i.e. ancho del pronotum a la altura de la dilatación supracoxal dividido por la longitud total de la metazona) del macho igual o superior a 0.63, en la hembra igual o superior a 0.63; metazona corta y ancha 2, Grupo Cayennensis
Medida de la forma del pronotum del macho y la hembra igual o menor a 0.43. Pronotum poco o muy elongado. Medida de la forma de la metazona (i.e. ancho del pronotum a la altura de la dilatación supracoxal dividido por la longitud total de la metazona) del macho igual o menor a 0.61, en la hembra igual o superior a 0.615 5
Grupo Cayennensis
2 Pronotum ensanchado en su extremo posterior, márgenes laterales sobresalientes; región central de la metazona con estrechamiento notorio (Fig. 47E–F) 3
Pronotum con márgenes laterals estrechándose gradualmente en la sección comprendida entre la dilatación supracoxal y el márgen posterior, sin o con muy poco estrechamiento central; márgenes laterales no sobresalientes y contenidos dentro del perfil del márgen más posterior (Fig. 47A–D). Parte distal de la apófisis faloide (afa) den esclerito dorsal derecho del macho (L4B) amplio en la base, estrechándose súbitamente para formar una punta aguda o roma; frecuentemente triangular (Fig. 51A–B) 4
3 Márgenes laterales de la prozona paralela o ligeramente divergentes anteriormente en ambos sexos (Fig. 47E–F). Región discoidal de las alas posteriores de ambos sexos de color naranja oxido, tornándose ahumada distalmente y hacia la región anal (Fig. 6A–B). Ápice de la apófisis faloide (afa) del esclerito dorsal derecho del macho (L4B) estrecho y digitiforme (Fig. 51C.1). Distribución en el Norte de Guyana y Norte de Brasil Liturgusa guyanensis La Greca, 1939
Márgenes laterales de la prozona de la hembra convergiendo anteriormente (Fig. 47G). Región discoidal de las alas posteriores de la hembra (unico sexo conocido) amarillo sobre el 90% más proximal y tornándose negro distalmente; la coloración amarilla se extiende hacia la porción anterior de la región anal, con una transición bien definida desde el amarillo hasta una condicioón más hialina, y con un tono mas ahumado hacia el márgen distal (Fig. 7A). Solo conocida para el Sur de Venezuela, cerca de la ”Sierra de la Neblina” Liturgusa neblina sp. n.
4 Mesotarso con primer segmento tan largo como los demás combinados. Distribuidos en el este de Sudamérica incluyendo Guyana Francesa, Guyana, Surinam y este de Brasil. Márgenes laterales del abdomen de la hembra con ligeras expansions tergales o sin estas Liturgusa cayennensis Saussure, 1869
Mesotarso con primer segmento más corto que los demás combinados. Distribución en el Oeste de Sudamérica, incluyendo Colombia, Ecuador, Perú y Sur de Venezuela. Márgenes laterales del abdomen de la hembra con expansions tergales notorias en forma de una punta orientada posteriormente Liturgusa lichenalis Gerstaecker, 1889
5 Medida de la forma del pronotum del macho igual o mayor a 0.360, de la hembra igual o mayor a 0.370 6, Grupo Maya
Medida de la forma del pronotum del macho igual o menor a 0.355, de la hembra igual o menor a 0.36; pronotum alargado y delgado 15, Grupo Cursor
Grupo Maya
6 Alas anteriores de la hembra 26–27 mm, del macho 19–20. Longitud del pronotum de la hembra 11–12 mm, del macho 9–10. Conocida solo para Colombia Liturgusa stiewei sp. n.
Alas anteriores de la hembra 13–22 mm (rango observado 13.35–21.41 mm), del macho 12–18 (rango observado 12.36–17.91 mm). Longitud del pronotum de la hembra 6–10 mm (rango observado 6.26–9.53 mm), del macho 5-8 mm (rango observado 5.16–7.35 mm) 7
7 Pronotum del macho y la hembra muy esculpido, mostrando las siguientes características: prozona con márgenes fuertemente convexos; dilatación supracoxal definida; región media de los márgenes laterales de la metazona paralelas o fuertemente convexa y dilatándose posteriormente; márgenes posteriores redondeados en los machos, en las hembras con las esquinas posteriores rectas, las que forman un angulo de 45 grados con respecto a la linea central del cuerpo (Fig. 47E–F). Distribuida solo en Centro America, con registros en Honduras, Guatemala, Belize y Panama Liturgusa zoae sp. n.
Pronotum del macho y la hembra no muy esculpido, mostrando las siguientes características: prozona con márgenes laterales casi paralelos o estrechándose anteriormente; dilatación supracoxal moderadamente definida; región media de los márgenes laterales de la metazona cóncavos o ligeramente dilatados; márgenes posteriores redondeados y con una emarginación central en ambos sexos (los machos de Liturgusa manausensis tienen las esquinas porteriores formando un angulo de 45 grados con respecto a la línea central del cuerpo (Figs 47H–T, 48C–D) 8
8 Relativamente pequeños. El producto de longitud de las alas anteriores multiplicado por la longitud del pronotum es igual a 63–68 en los machos, y 88–91 en las hembras 9
Relativamente grandes. El producto de longitud de las alas anteriores multiplicado por la longitud del pronotum es igual a 73–132 en los machos, y 97–187 en las hembras 10
9 Fémures anteriores de la hembra con espina genicular posteroventral posicionada proximalmente al origen del lóbulo genicular. Rango de la siguiente métrica relativa: (longitud del pronotum / ancho menor del pronotum) × ancho de la cabeza, igual a 16–17 en los machos Liturgusa bororum sp. n.
Fémures anteriores de la hembra con espina genicular posteroventral posicionada distalmente al origen lóbulo genicular. Rango de la siguiente métrica relativa: (longitud del pronotum / ancho menor del pronotum) × ancho de la cabeza, aproximadamente 14 en los machos Liturgusa cura sp. n.
10 Alas posteriores del macho y la hembra con una coloracion amarilla, naranja u oxido sobre al menos el tercio proximal de la region discoidal, tornandose negra distalmente (Figs 10A–B, 16A–B) 11
Región discoidal de las alas posteriores del macho y la hembra de color negro u oscuramente opacada, con a lo mucho el margen anterior del márgen costal de un color mas pálido o algo rojizo (Figs 11A–B, 12A, 13A–B, 17A–B) 12
11 Área discoidal de las alas posteriores del macho y la hembra naranja o amarillo en los ¾ proximales, rápidamente tornándose negro distalmente; región anal mayormente negra u oscuramente opacada, con el tercio proximal coloreado como la región discoidal (Fig. 10A–B). La métrica longitud del fémur anterior / longitud de las alas anteriores en los machos es igual o mayor a 0.45 (rango observado 0.45–0.48), e igual o mayor a 0.47 en las hembras (rango observado 0.47–0.52) Esclerito ventral izquierdo (L4A) sin proceso distal (pda) y suavemente redondeado. Distribución en Centro América, con registros de Panamá, Costa Rica y Nicaragua Liturgusa fossetti sp. n.
Área discoidal de las alas posteriores del macho y la hembra naranja o rojo oxido en los 1/3 al ½ proximales, gradualmente tornandose negro hacia distalmente. Región anal uniformemente opaca, negra o ahumada (Fig. 16A–B). La métrica longitud del fémur anterior / longitud de las alas anteriores en los machos igual o menor a 0.44 (rango observado 0.4–0.44), e igual o menor a 0.45 en las hembras (rango observado 0.4–0.45). Esclerito ventral izquierdo (L4A) con proceso distal (pda) (Fig. 51J.1–J.6) Liturgusa tessae sp. n.
Diseccion de genitalia masculina es necessaria para la identificacion de Liturgusa manausensis, Liturgusa kirtlandi, Liturgusa trinidadensis, and Liturgusa maya.
12 Métrica de la forma del pronotum del macho 0.59 (métrica = [longitud de la prozona/longitud del pronotum] × ancho del pronotum a la altura de la dilatación supracoxal]. Apófisis faloide (afa) del esclerito dorsal izquierdo (L4B) con una punta larga y aguda; ventral plate (pia) y ventral process (pva) del falómero derecho (R1) grandes (Fig. 51H.1). Especie pequeña con coloración de las alas anteriores asimétrica y con un contraste muy notorio entre las regiones oscurecidas y las más claras. Especie conocida solo por los machos, registros solo para Manaus, Brasil Liturgusa manausensis sp. n.
Métrica de la forma del pronotum del macho 0.63 o mayor (rango observado 0.63-0.86), en las hembras 0.77 o mayor (rango observado 0.77-1.12). Apófisis faloide (afa) del esclerito dorsal izquierdo (L4B) con una punta corta y aguda; ventral plate (pia) y ventral process (pva) del falómero derecho (R1) pequeños (Figs 51G.1–G.4, 51I.1–I4, 52A.1–A.5) 13
13 Proceso apical (paa) del esclerito dorsal izquierdo (L4B) plano, en forma de podio (Fig. 51G.1–G.4). Especie relativamente grande, pronotum del macho entre 6.95–7.35 (promedio 7.18), y hembras entre 8.16–8.7 (promedio 8.43). Registros solo para Bolivia central Liturgusa kirtlandi sp. n.
Proceso apical (paa) del esclerito dorsal izquierdo (L4B) uniformemente redondeado (Figs 51I.1–I4, 52A.1–A.5). Tamaño relativamente pequeño, pronotum del macho entre 5.61–7.39 (promedio 6.25), y hembras 6.55–8.99 (promedio 7.42) (solo se observo un macho y dos hembras de Liturgusa maya con medidas del pronotum comparables a Liturgusa kirtlandi) 14
14 Esquinas posterolaterales de la metazona con numerosos tubérculos en ambos sexos. Antenas pálidas o gradualmente tornándose pardo en ambos sexos. Proceso distal (pda) del esclerito ventral izquierdo (L4A) centralmente con un pliegue esclerotizado o una ligera proyección (Fig. 52A.1–A.5). Registros solo para Trinidad Liturgusa trinidadensis sp. n.
Sin tubérculos en el pronotum de ambos sexos. Antenas tornándose rápidamente negras distalmente en ambos sexos. Proceso distal (pda) del esclerito ventral izquierdo (L4A) centralmente con un proceso fuerte bien definido, el cual forma un ángulo de ca. 30 grados con el eje central del hypophallus, el ápice es redondeado (Fig. 51I.1–I4). Registros solo para Trinidad Liturgusa maya Saussure & Zehntner, 1894
15 Longitud de las alas del macho igual o menor a 13.2. Métrica longitud de las alas anteriores × longitud del pronotum del macho igual o menos a 92, igual o menor a 125 en las hembras. Esclerito ventral izquierdo (L4A) con el ápice liso y redondeado, carente de proceso distal (pda) (Fig. 52C.1–E.1) 16, Grupo Cursor A
Longitud de las alas del macho igual o menor a 13.4. Métrica longitud de las alas anteriores × longitud del pronotum del macho igual o mayor a 95, igual o mayor a 130 en las hembras. Esclerito ventral izquierdo (L4A) con un proceso distal (pda) o un borde aserrado (Fig. 52F.1–K.1) 18, Grupo Cursor B
Grupo Cursor A
16 Esclerito supra-anal del macho y la hembra fuertemente transversal, aproximadamente 3 veces tan ancho como largo. Alas anteriores sin una marca blanca fuertemete pronunciada cerca del extreme proximal (Fig. 19A–B). Apófisis faloide (afa) del esclerito dorsal derecho (L4B) digitiforme, con el ápice en forma de un bulbo ligeramente ensanchado (Fig. 52C.1–C.3). Distribución solo en Centro América, con registros para Nicaragua, Costa Rica y Panamá Liturgusa cursor Rehn, 1950
Esclerito supra-anal del macho y la hembra aproximadamente 2 veces tan ancho como largo, con ápice redondeado. Alas anteriores con una marca blanca fuertemente pronunciada sobre el 1/5 proximal y sobre la base de las venas media y cubital (Figs 20A–B, 22A–B). Apófisis faloide (afa) del esclerito dorsal izquierdo (L4B) triangular, terminando en punta aguda (Fig. 52D.1–E.1) 17
17 Superficie anterior de la procoxa con una banda media de color negro en la mitad proximal, así como también una marca negra medial hacia el ápice. Margen posterior del fémur anterior con una espina pequeña entre la espina posteroventral más distal y la espina genicular. Espina genicular posicionada proximalmente al inicio del lóbulo genicular. Distribución en el norte de Sudamérica, con registros para Guyana Francesa, pero posiblemente distribuida también en Surinam y norte de Brasil Liturgusa milleri sp. n.
Superficie anterior de la procoxa pálida, sin marcas negras. Margen posterior del fémur anterior sin una espina definida entre la espina posteroventral más distal y la espina genicular. Espina genicular posicionada distalmente al inicio del lóbulo genicular. Endémica de la isla Dominica Liturgusa dominica sp. n.
Grupo Cursor B
18 Parte proximal del área discoidal de las alas posteriores de color rojo, naranja u óxido, tornándose negro distalmente (Figs 26A–B, 29A). Ala anterior del macho y la hembra frecuentemente de coloración asimétrica, cuando esto ocurre, una de las alas tiene un tono rojizo 19
Área discoidal de las alas posteriores uniformemente oscurecida o ahumada (Figs 7B, 23A–B, 24A–B, 25B, 27A–B). Alas anteriores del macho y la hembra con o sin coloración asimétrica, pero cuando esta ocurre, una es oscura, negra o ahumada 20
19 Espinas posteroventrales del fémur anterior largas, tanto en machos como en las hembras, siendo la más proximal aproximadamente 0.5 y 1 mm, respectivamente (longitud del fémur anterior ~14 y ~8.5 veces la longitud de la espina posteroventral más proximal, respectivamente). Apófisis faloide (afa) del esclerito dorsal izquierdo (L4B) delgado, como una aguja; ápice del esclerito ventral izquierdo (L4A) con una proyección aguda y ligeramente curvada, aunque puede estar reducida a una distal process (pda) más corta y de ápice algo puntiagudo (Fig. 52I.1) Liturgusa krattorum sp. n.
Espinas posteroventrales del fémur anterior cortas, tanto en machos como en las hembras, siendo la más proximal aproximadamente 0.35 y 0.5 mm, respectivamente (longitud del fémur anterior 18 y 15.5 veces la longitud de la espina posteroventral más proximal, respectivamente). Apófisis faloide (afa) del esclerito dorsal izquierdo (L4B) en forma de un triangulo amplio, terminando en una punta aguda; ápice del esclerito ventral izquierdo (L4A) con una distal process (pda) corta y roma (Fig. 52K.1) Liturgusa purus sp. n.
20 Longitud del pronotum de los machos menor a 7.1, el de las hembras menor a 8 Liturgusa actuosa Rehn, 1950
Longitud del pronotum de los machos mayor a 7.1, el de las hembras mayor a 8.5 21
21 Longitud de las alas anteriores como máximo 1.80 veces más largo que la longitud total del pronotum (rango observado 1.67–1.80). Ala posterior del macho y la hembra con una emarginación entre las áreas discoidal y anal, poco o muy poco marcada (Figs 7A, 25B) 22
Longitud de las alas anteriores como minimo 1.83 veces más largo que la longitud total del pronotum (rango observado 1.83–2.2). Ala posterior del macho y la hembra con una obvia emarginación entre las áreas discoidal y anal (Figs 25A–B, 27A–B) 23
22 La segunda espina posteroventral más proximal de las tibias anteriores de la hembra, aproximadamente 1.4 veces la longitud de la tercera espina. Terga abdominal de la hembra con pequeñas expansiones en las esquinas latero-posteriores. Distribución en el norte de Venezuela, con un registro en el norte de Guyana Liturgusa cameroni sp. n.
La segunda y tercera espina posteroventral mas proximales de las tibias anteriores de la hembra, casi de la misma longitud, con la segunda siendo solo ligeramente más larga en la mayoría de especimens examinados (no más de 1.2 veces la longitud de la tercera espina). Terga abdominal de la hembra con expansiones pronunciadas en las esquinas latero-posteriores de aproximadamente 0.15 y 0.21 mm de longitud. Distribución en la cuenca del Amazonas Liturgusa maroni sp. n.
23 Primer tarsómero de la pata posterior de la hembra 3.5–8 veces más largo que la espina apical dorsal de la metatibia (dicha espina es relativamente larga). Proceso distal (pda) del esclerito ventral izquierdo (L4A) en punta, uno de sus lados es aserrado y fuertemente esclerotizado; apofisis faloide (afa) del esclerito dorsal izquierdo (L4B) con un apice amplio y triangular (Fig. 52J.1–J.2). Liturgusa nubeculosa Gerstaecker, 1889
Primer tarsómero de la pata posterior de la hembra 10-12 veces más largo que la espina apical dorsal de la metatibia (dicha espina es relativamente corta). Proceso distal (pda) del esclerito ventral izquierdo (L4A) en punta redondeada; apofisis faloide (afa) del esclerito dorsal izquierdo (L4B) con un apice Delgado y en punta (Fig. 52G.1–G.2) Liturgusa algorei sp. n.
Cayennensis Group
Liturgusa cayennensis Saussure, 1869

http://species-id.net/wiki/Liturgusa cayennensis

Liturgousa cayennensis: Saussure 1869: 62; Brauer 1870: 92; Saussure 1871b: 101–102; Saussure and Zehntner 1894: 159–160; Westwood 1889: 5, 50; Scudder 1901: 159, 407; Kirby 1904: 271; Chopard 1911: 323; Chopard 1916: 164; Hebard 1919a: 31; Hebard 1924: 131; Hebard 1929: 399; Hebard 1933: 29; Rehn 1935: 198; Beebe et al. 1952: 246; Cerdá 1996: 75–76; Roy and Cuche 2008: 8, 21.
Liturgusa cayennensis: Hebard 1922: 337; Giglio-Tos 1927: 293; Beier 1935: 11; Jantsch 1991: 125; Terra 1995: 53; Jantsch 1999: 47–48; Salazar E. 2000: 67; Lombardo and Agabiti 2001: 90, 96; Salazar E. 2002: 124; Ehrmann 2002: 206; Agudelo 2004: 55, Table 3.1; Agudelo 2005: 3; Otte and Spearman 2005: 132; Agudelo et al. 2007: 116, 141.
Liturgousa cayennesis: Bruner 1906: 143.
Type.

Holotype Female. Muséum d’Histoire naturelle, Geneva, Switzerland

Type locality.

Cayenne. (French Guiana)

Material examined.

Liturgusa cayennensis Saussure, 1869.

Sex Type Country Label Latitude Longitude Code
Male nontype Guyana B.G. 5-1-1923, W. Bank, Dem. R. AMNH 030
Female nontype Guyana Kartabo, Bartica District, British Guiana, 1921 6.242050, -59.306552 ANSP 046
4 Females nontype Guyana Bartica, British Guiana, H.S. Parish, 1.20.1912 6.405831, -58.625444 ANSP 048-51
Male nontype Suriname Ongelijk, Para, R. Surinam, May‚ 27 ANSP 054
Male nontype French Guiana Guyane Francse, Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, Mai ANSP 055
Female nontype Guyana Guyane, 1907-247. BMNH 007
Male nontype Brazil Mato Grosso, Serra do Roncador, 264 km N. of Xavantina, near base camp, 1967-9, I.R. Bishop, RS/RGS exp. B.M. 1981-312 -12.456161, -52.054393 BMNH 076
Male nontype Guyana New River, 750 ft., 10-20.III.1938, C.A. Hudson 2.543686, -57.583411 BMNH 086
Male nontype French Guiana Kaw Mountain Res., Amazonas Lodge, 4°32'57.8"N, 52°12'49.7"W, 8-19 Feb 2005, Coll: K.B. Miller 4.549389, -52.213806 GSMC000262
Nymph nontype French Guiana Maripasoula, Saul. Campsite Chez Fred‘s, 03°37'42.0"N, 53°12'40.0"WEle:26m, Manicured landscape surrounded by Primary tropical rainforest, Hand collected from leaf litter samples, Dec. 14-15, 2004, Coll: J. Huff 3.628333, -53.211111 GSMC003069
Female nontype French Guiana St-Jean du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Fevrier 5.487038, -54.008462 MNHN 001
Female nontype French Guiana Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, June MNHN 002
Male nontype French Guiana Cayenne 4.914356, -52.301354 MNHN 005
Female nontype Brazil Amazonas Reserva Biologica do Cuieiras, 50 km. N. Manaus, 15-IV-AU 15-V-1981, M. Descamps, Bresil -2.599556, -60.210631 MNHN 0059
Male nontype French Guiana Cayenne 4.914356, -52.301354 MNHN 007
Female nontype French Guiana Saut Dalles, 24-VIII-1992, P. Peters, Guyane francaise 3.291071, -53.835810 MNHN 0075
Female nontype French Guiana Kourou, Guyane Franc, Coll: A. Bonhoure, 1909, Avril 5.163511, -52.656581 MNHN 011
Female nontype French Guiana St-Jean du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Decembre 5.487038, -54.008462 MNHN 012
Female nontype French Guiana Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, Novembre MNHN 013
Female nontype French Guiana St-Jean du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Fevrier 5.487038, -54.008462 MNHN 014
Male nontype French Guiana Petit Saut, 8-II-1994, P.E. Rouland 5.069416, -53.047566 MNHN 034
Female nontype French Guiana Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, Novembre MNHN 050
Male nontype French Guiana Sinnamary, VII-1977, Guyane, M. Descamps rec. 5.370512, -52.960320 MNHN 065
Male nontype French Guiana Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, Mai MNHN 066
Male nontype French Guiana St-Jean du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Janvier 5.487038, -54.008462 MNHN 070
Male nontype French Guiana St-Jean du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Janvier 5.487038, -54.008462 MNHN 071
Female nontype French Guiana Pied Saut Parare, 12-VIII-1977, Guyane, M. Descamps Rec. 4.046724, -52.698087 MNHN 083
Female nontype Guyana Mazaruni-Potaro: Waratuk Falls, Potaro R, 1 rainforest 300 ft, 1 Oct 1990, ROM 905028, LD Coote 5.272097, -59.397117 ROM 003
Male nontype Suriname ex. coll. H. Dohrn SDEI 002
Female nontype French Guiana St-Laurent du Maroni, Collection Wm Schaus 5.487038, -54.008462 USNM 033: USNM ENT 00873000
Male nontype Brazil Brasilien Amazonas, Parintins, Dr. F. Knutsen -2.653184, -56.732240 ZMHB 001
Female nontype Brazil Amazonas, oberer Solimoes., Rio Tomantins 9.26, Eing.Nr.33. 1937 ZMUH 005
Male nontype French Guiana unknown MNHN 074
Taxonomic history.

The oldest species described within the genus is also the type species for Liturgusa, designated by Kirby in 1904. Although the species was included in numerous taxonomic works, the species was likely confused with numerous others since some of these works focus on regions where Liturgusa cayennensis does not range. Therefore, it is impossible to tell what species they had misidentified as Liturgusa cayennensis. It was Rehn (1935) that first started to notice that Liturgusa cayennensis was more geographically restricted and that many previous records for the species in Central America were not correct. He noticed that it was being confused with Liturgusa maya most often. For Liturgusa, identification has always been a problem and most previous records of Liturgusa cayennensis should be viewed carefully and probably not included within distributional studies.

Diagnosis.

A short and stocky species with shortened, rounded forewings, a short and broad pronotum, and a broad abdomen. The forewings are usually asymmetrical in coloration, the darker being rust colored. Most similar to Liturgusa lichenalis, but lacking the prominent posterolateral tergal projections. Both Liturgusa guyana and Liturgusa neblina are similar, but both have a more pronounced constriction in the metazone of the pronotum. Finally, Liturgusa lichenalis is restricted to the western Amazon basin while Liturgusa cayennensis is found in central and eastern regions of the Amazon basin.

Description.

Male. (Fig. 3A) N=9: Body length 19.22–22.58 (20.93); forewing length 12.49–14.98 (13.61); hindwing length 10.26–11.37 (10.63); pronotum length 5.31–6.01 (5.60); prozone length 1.68–2.00 (1.79); pronotum width 2.38–2.70 (2.53); pronotum narrow width 1.77–1.98 (1.85); head width 4.84–5.39 (5.03); head vertex to clypeus 1.83–2.07 (1.96); frons width 1.67–1.87 (1.79); frons height 0.64–0.82 (0.72); prothoracic femur length 5.52–6.57 (6.00); mesothoracic femur length 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.

Figure 3.

Liturgusa cayennensis Saussure, 1869, dorsal habitus: A male from French Guiana (MNHN 034) B female from Guyana (ANSP 049).

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 supra-coxal 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.

Redescription.

Female. (Figs 3B, 4A) N=13: Body length 14.64–32.95 (28.65); forewing length 17.01–20.50 (19.21); hindwing length 14.50–16.43 (15.64); pronotum length 7.03–8.11 (7.70); prozone length 2.23–2.63 (2.46); pronotum width 3.32–4.14 (3.63); pronotum narrow width 2.30–3.09 (2.60); head width 6.40–7.13 (6.81); head vertex to clypeus 2.74–3.10 (2.93); frons width 2.46–2.92 (2.73); frons height 0.98–1.14 (1.07); prothoracic femur length 7.46–9.15 (8.29); mesothoracic femur length 8.41–9.91 (9.12); mesothoracic tibia length 5.34–7.81 (7.05); mesothoracic tarsus length 5.10–7.06 (6.46); metathoracic femur length 8.62–9.78 (9.18); metathoracic tibia length 9.45–10.86 (9.97); metathoracic tarsus length 8.37–9.57 (9.13); pronotal elongation measure 0.31–0.33 (0.32); pronotal shape measure 0.45–0.52 (0.47); head shape measure 0.42–0.44 (0.43); frons shape measure 0.36–0.43 (0.39); anteroventral femoral spine count 15–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 4.

Liturgusa cayennensis Saussure, 1869, and Liturgusa lichenalis Gerstaecker, 1889, dorsal habitus of types and labels. Liturgusa cayennensis: A holotype female (MHNG) B labels. Liturgusa lichenalis: C holotype female (EMAU) D labels.

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.

Meso- and Metathoracic Legs: Femora with expanded ventral (posterior) carina, almost a lamellar expansion; dorsal (anterior) carina very pronounced.

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. Supra-anal plate slightly transverse, and evenly rounded lobe.

Liturgusa lichenalis Gerstaecker, 1889

http://species-id.net/wiki/Liturgusa_lichenalis

Liturgusa lichenalis: Gerstaecker 1889: 52–53; Bertkau 1889: 87; Terra 1995: 54; Ehrmann 2002: 207; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116.
Liturgousa lichenalis: Westwood 1889: 5, 51; Kirby 1904: 271; Caudell 1918: 5; Rehn 1935: 199; Rehn 1954: 179, pl. 1, fig. 1.
Liturgusa annulipes (partim): Giglio-Tos 1927: 294; Beier 1935: 11; Jantsch 1999: 48.
Type.

Holotype Female. Ernst-Moritz-Arndt-Universität Greifswald, Germany.

Type locality.

Peru: Jurimaguas (Lat. -5.900438, Long. -76.125298).

Material examined.

Liturgusa lichenalis Gerstaecker, 1889

Sex Type Country Label Latitude Longitude Code
Female Holotype Peru Jurimaguas -5.900438, -76.125298 EMAU
Male nontype Peru Monson Valley, Tingo Maria, XII-11-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 001
Male nontype Peru Monson Valley, Tingo Maria, XII-11-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 005
Male nontype Peru Monson Valley, Tingo Maria, XII-11-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 010
Male nontype Peru Chanchamayo, 04-08-05, E. Cueva -11.135912, -75.348800 MEKRB 010
Female nontype Peru Dept. Loreto Colonia, Amont Conflt. Rios Zumun & Yahuasyacu, 20-V-20-VI-1978, M. Descamps rec MNHN 055
Female nontype Colombia Dept. Amazonas, Rio Igara Parana, 30 km aval La Chorrera, VI - VII 1974, M. Descamps rec. -1.197385, -72.937475 MNHN 081
Male nontype Peru Loreto - Maynas, Picurovacu, apres Sta Clotilde, N. Iquitos, 130 mts - 10.II.2010, S03.37.04 - W 73.15.44, coll. M.Dottax -3.617778, -73.262222 MNHN 100
Female nontype Ecuador Pr. Sucumbios. Coca/L. Agrio, S.P. de los Cofanes, 415 m. 22/23 IX 1997, 77°52'W, 0°8'S, Amedegnato/Poulain rec. -0.133333, -77.866667 MNHN 203
Male nontype Venezuela Culebra, N. Duida Territ. Amazonas, April 7-16, 1950, J. Maldonado Capriles Coll. 3.198590, -65.555990 USNM 016; USNM ENT 00873001
Female nontype Peru 11°3'S, 75°17'W, X.08, GREENW, C. Schunke -11.050000, -75.283333 USNM 059; USNM ENT 00873002
Female nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 02 Mar 84, 03/02 -12.833333, -69.283333 USNM 071; USNM ENT 00873039
Female nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 01 Mar 82, 01/034/02 -12.833333, -69.283333 USNM 072; USNM ENT 00873040
Female nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 10Sept84, 02/02/055 -12.833333, -69.283333 USNM 073; USNM ENT 00873041
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 07Nov83, 01/02/71 -12.833333, -69.283333 USNM 074; USNM ENT 00873042
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 10Sept84, 02/03/114 -12.833333, -69.283333 USNM 075; USNM ENT 00873043
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 30Apr84, 03/03/089 -12.833333, -69.283333 USNM 076; USNM ENT 00873044
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 02Mar84, 03/02/056 -12.833333, -69.283333 USNM 077; USNM ENT 00873045
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 14Sept84, 01/03/120 -12.833333, -69.283333 USNM 078; USNM ENT 00873046
Male nontype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 04Mar84 -12.833333, -69.283333 USNM 079; USNM ENT 00873047
Male nontype Peru Madre de Dios, Rio Tambopata Res., 12°50'S, 69°17'W; Coll: G.J. Svenson 2005 -12.833333, -69.283333 GSMC000259
Male nontype Ecuador Pastaza; Ashuara, Rio Macuma, 10km from Rio Morona, 300m, VII:7-16:1971, leg. B. Malkin -2.753512, -77.444899 FMNH 001
Male nontype Peru Loreto; Yagua, Indian village, head-waters Rio Loreto-Yacu, IV:29-V:1:1970, leg. B. Malkin -3.894566, -71.971608 FMNH 002
Female nontype Peru Loreto; Ucayali R., Yarina Cocha, II-27-1956, leg. Peter Hocking -5.509011, -74.377603 FMNH 007
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 Liturgusa lichenalis as the synonym of Fuga annulipes, but this treatment is erroneous as it is apparent that no previous work had an accurate concept of Fuga annulipes. Rehn (1935) corrected this erroneous synonym by comparing the types of both species.

Diagnosis.

Almost identical to Liturgusa 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 Liturgusa cayennensis and have posterolateral tergal projections in the distal half. Another distinguishing external character for Liturgusa lichenalis comparison to Liturgusa 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 Liturgusa cayennensis.

Description.

Male. (Fig. 5A) N=12: Body length 18.59–24.79 (21.45); forewing length 12.62–16.54 (14.17); hindwing length 10.27–11.75 (11.01); pronotum length 5.03–6.59 (5.74); prozone length 1.55–2.07 (1.78); pronotum width 2.33–3.18 (2.68); pronotum narrow width 1.78–2.47 (2.07); head width 4.54–5.75 (5.03); head vertex to clypeus 1.81–2.03 (1.91); frons width 1.66–1.93 (1.78); frons height 0.59–0.75 (0.67); prothoracic femur length 5.60–7.14 (6.19); mesothoracic femur length 6.37–8.50 (7.38); mesothoracic tibia length 4.78–6.27 (5.52); mesothoracic tarsus length 4.77–6.25 (5.32); metathoracic femur length 6.71–8.92 (7.55); metathoracic tibia length 6.82–8.68 (7.64); metathoracic tarsus length 6.41–8.40 (7.34); pronotal elongation measure 0.30–0.33 (0.31); pronotal shape measure 0.44–0.49 (0.47); head shape measure 0.38–0.40 (0.39); frons shape measure 0.35–0.40 (0.38); anteroventral femoral spine count 13–17 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 5.

Liturgusa lichenalis Gerstaecker, 1889, dorsal habitus: A male from Tingo Maria, Peru (CAS 010) B female from Ecuador (MNHN 203).

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.

Meso- and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina pronounced. Mesotarsi with first segment shorter than 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 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.

Redescription.

Female. (Figs 4C, 5B) N=8: Body length 25.68–32.17 (28.36); forewing length 16.25–19.99 (18.46); hindwing length 13.46–16.50 (14.96); pronotum length 6.68–8.26 (7.48); prozone length 2.12–2.64 (2.36); pronotum width 3.29–3.99 (3.59); pronotum narrow width 2.49–2.82 (2.71); head width 5.96–7.01 (6.48); head vertex to clypeus 2.55–3.05 (2.81); frons width 2.36–2.87 (2.62); frons height 0.93–1.13 (1.01); prothoracic femur length 7.40–9.15 (8.11); mesothoracic femur length 8.30–9.76 (8.85); mesothoracic tibia length 6.39–8.13 (7.04); mesothoracic tarsus length 5.67–6.75 (6.19); metathoracic femur length 8.18–9.91 (8.75); metathoracic tibia length 8.59–11.05 (9.60); metathoracic tarsus length 7.73–9.35 (8.51); pronotal elongation measure 0.31–0.33 (0.32); pronotal shape measure 0.47–0.51 (0.48); head shape measure 0.42–0.44 (0.43); frons shape measure 0.36–0.41 (0.39); anteroventral femoral spine count 15–16 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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. Supra-anal plate slightly transverse, margins tapering gradually to a rounded terminus.

Liturgusa guyanensis La Greca, 1939

http://species-id.net/wiki/Liturgusa_guyanensis

Liturgusa guyanensis: La Greca 1939: 2–5, fig. 1; Terra 1995: 54; Ehrmann 2002: 207; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116.
Liturgusa guyannensis: Jantsch 1999: 48.
Type.

LOST, but listed repository Natural History Museum Zoological Section 'La Specola', University of Florence.

Original type locality.

Guyana, Babooncamp (Demerara), October 1931.

Neotype.

Male. California Academy of Sciences, San Francisco, CA, USA.

Neotype locality.

Guyana: Iwokrama, Forest Research Station, 1 km N. Kurupukari, 14-19 January 1996, canopy fog sample of Mora tree, W. Tschinkel coll. (Lat. 4.672093, Long. -58.685606).

Material examined.

Liturgusa guyanensis La Greca, 1939.

Sex Type Country Label Latitude Longitude Code
Male Neotype Guyana Iwokrama, Forest Research Station, 1 km N. Kurupukari, 14-19 January 1996, canopy fog sample of Mora tree, W. Tschinkel coll. 4.672093, -58.685606 CAS 021
Female nontype Brazil Manaos, II-III-43, Thomas Gilliard -3.003866, -59.961947 AMNH 027
Male nontype Guyana Iwokrama, Forest Research Station, 1 km N. Kurupukari, 14-19 January 1996, canopy fog sample of Mora tree, W. Tschinkel coll. 4.672093, -58.685606 CAS 017
Nymph nontype Brazil Amazonas, Rio Taruma Mirim, 20 km nw Manaus, 02 Mar 1979, 02°53'S, 060°07'W, Montgomery, Erwin, Schimmel, Krischik, Date, Bacon colls., Black water innundation forest canopy fogged with Pyrethrum, Sample # 48. -2.883333, -60.116667 USNM 010; USNM ENT 00873003
Taxonomic history.

Described in 1939 by Marcello La Greca, Liturgusa 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 Liturgusa 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 Liturgusa charpentieri nor Liturgusa 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.

Diagnosis.

Similar to Liturgusa cayennensis and living in sympatry with at least some populations of the species, Liturgusa 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 Liturgusa cayennensis and Liturgusa lichenalis).

Description.

Male. (Fig. 6A) N=1: Body length 23.28; forewing length 15.42; hindwing length 12.31; pronotum length 6.19; prozone length 1.85; pronotum width 2.83; pronotum narrow width 1.92; head width 5.46; head vertex to clypeus 2.20; frons width 2.07; frons height 0.82; prothoracic femur length 6.41; mesothoracic femur length 7.69; metathoracic femur length 7.57; pronotal elongation measure 0.30; pronotal shape measure 0.46; head shape measure 0.40; frons shape measure 0.39; anteroventral femoral spine count 14; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 6.

Liturgusa guyanensis La Greca, 1939, dorsal habitus: A neotype male from Iwokrama, Guyana (CAS 021) B female from Manaus, Brazil (AMNH 027).

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 juxta-ocular 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.

Redescription.

Female. (Fig. 6B) N=1: Body length 26.83; forewing length 18.06; hindwing length 14.03; pronotum length 7.10; prozone length 2.09; pronotum width 3.50; pronotum narrow width 2.32; head width 6.63; head vertex to clypeus 2.97; frons width 2.79; frons height 1.11; prothoracic femur length 7.61; mesothoracic femur length 8.01; mesothoracic tibia length 5.92; mesothoracic tarsus length 5.80; pronotal elongation measure 0.29; pronotal shape measure 0.49; head shape measure 0.45; frons shape measure 0.40; anteroventral femoral spine count 14; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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. Supra-anal plate slightly transverse, and evenly rounded lobe.

Type.

Holotype Female, pinned. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Type Locality.

Venezuela: T.F. Amaz., Cerro de la Neblina, Basecamp, 140 m, 0°50'N, 66°10'W, 19 February 1985, Pyrethrin fogging of vine tangle: canopy of floodplain forest along Rio Baria; R. Cocroft & W. Steiner. (Lat. 0.833333, Long. -66.166667).

Material examined.

Liturgusa neblina sp. n.

Sex Type Country Label Latitude Longitude Code
Female Holotype Venezuela T.F. Amaz., Cerro de la Neblina, Basecamp, 140 m, 0°50'N, 66°10'W, 19 February 1985, Pyrethrin fogging of vine tangle: canopy of flood plain forest aling Rio Baria; R. Cocroft & W. Steiner. 0.833333, -66.166667 USNM 006; USNM ENT 00873998
Diagnosis.

Most similar in general appearance to Liturgusa cayennensis and Liturgusa lichenalis. The type locality overlaps with the range of Liturgusa lichenalis, but Liturgusa 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.

Description.

Female. (Fig. 7A) N=1: Body length 29.75; forewing length 19.54; hindwing length 16.35; pronotum length 8.15; prozone length 2.46; pronotum width 3.68; pronotum narrow width 2.46; head width 6.90; head vertex to clypeus 2.88; frons width 2.78; frons height 1.10; prothoracic femur length 8.45; mesothoracic femur length 9.50; mesothoracic tibia length 7.50; mesothoracic tarsus length 6.48; metathoracic femur length 9.42; metathoracic tibia length 10.72; metathoracic tarsus length 9.13; pronotal elongation measure 0.30; pronotal shape measure 0.45; head shape measure 0.42; frons shape measure 0.41; anteroventral femoral spine count 14; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 7.

Liturgusa, dorsal habitus: A Liturgusa neblina sp. n., holotype female from Cerro de la Neblina, Venezuela (USNM 006) B Liturgusa maroni sp. n., holotype female from St. Laurent du Maroni, French Guiana (MNHN 019).

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.

Meso- and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina pronounced. Mesotarsi with first segment shorter than the remaining segments combined.

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, Liturgusa 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.

Maya Group
Type.

Holotype Male, pinned. Muséum national d’Histoire naturelle, Paris, France.

Type locality.

Peru: Loreto, Brillo Nuevo, Reg. de l‘Ampiyacu, en av. confl. des rios Zumun et Yahuasyacu, L. Desutter Rec.; Paro.Q; Sans (5A. Apres Aband.) Chasse Jour 9-XI-1985 (Lat. -3.335174, Long. -71.816805).

Material examined.

Liturgusa bororum sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Peru Loreto, Brillo Nuevo, Reg. de l‘Ampiyacu, en av. confl. des rios Zumun et Yahuasyacu, L. Desutter Rec.; Paro.Q; Sans (5A. Apres Aband.) Chasse Jour 9-XI-1985 -3.335174, -71.816805 MNHN 038
Female Allotype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004008
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson - Tissue 005 -3.620960, -72.247440 GSMC004052
Female Paratype Brazil Benjamin Constant, IX 1979, AM, B. Silva rec., C. Seabia leg. -4.383010, -70.042251 MNHN 051
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 Liturgusa algorei, Liturgusa krattorum, and Liturgusa maya at the Madre Selva Biological Research Station in the Loreto Province, Peru.

Diagnosis.

The second smallest species in Liturgusa, second only to Liturgusa cura, Liturgusa bororum can be distinguished from Liturgusa cura easily by the presence of dark pigmentation across the ventral surface of the prothoracic femora in males and two markings in females; males of Liturgusa cura have only two small marks and females are entirely devoid of markings. Liturgusa bororum is only known from northern Peru and just across the border in Brazil. Can be distinguished from Liturgusa manausensis by its evenly rounded posterior margins of the metazone.

Description.

Male. (Fig. 8A) N=2: Body length 19.61–20.30 (19.95); forewing length 12.68–13.63 (13.15); hindwing length 9.88; pronotum length 5.35–5.64 (5.49); prozone length 1.58–1.61 (1.59); pronotum width 2.11–2.25 (2.18); pronotum narrow width 1.46–1.61 (1.53); head width 4.51–4.55 (4.53); head vertex to clypeus 1.80; frons width 1.59–1.71 (1.65); frons height 0.53–0.67 (0.60); prothoracic femur length 5.54–5.99 (5.77); mesothoracic femur length 7.43–7.71 (7.57); mesothoracic tibia length 5.81–6.23 (6.02); mesothoracic tarsus length 5.11; metathoracic femur length 7.27–7.59 (7.43); metathoracic tibia length 7.75–8.54 (8.15); metathoracic tarsus length 8.07; pronotal elongation measure 0.29; pronotal shape measure 0.40; head shape measure 0.40; frons shape measure 0.31–0.42 (0.36); anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 8.

Liturgusa bororum sp. n., dorsal habitus: A holotype male from Brillo Nuevo, Peru (MNHN 038) B allotype female from Loreto, Peru (CLEV GSMC004008).

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 c-shaped; 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.

Female. (Fig. 8B) N=2: Body length 24.01–24.69 (24.35); forewing length 14.54–15.55 (15.04); hindwing length 11.29–11.83 (11.56); pronotum length 6.26–6.43 (6.34); prozone length 1.90–1.96 (1.93); pronotum width 2.47–2.66 (2.56); pronotum narrow width 1.72–1.82 (1.77); head width 5.18–5.58 (5.38); head vertex to clypeus 2.19–2.43 (2.31); frons width 2.01–2.25 (2.13); frons height 0.82–0.90 (0.86); prothoracic femur length 6.33–6.58 (6.46); mesothoracic femur length 8.20–8.21 (8.20); mesothoracic tibia length 6.49–6.53 (6.51); mesothoracic tarsus length 6.01–6.24 (6.13); metathoracic femur length 8.26–8.54 (8.40); metathoracic tibia length 8.69–8.81 (8.75); metathoracic tarsus length 8.81–9.16 (8.98); pronotal elongation measure 0.30; pronotal shape measure 0.40–0.41 (0.40); head shape measure 0.42–0.44 (0.43); frons shape measure 0.40–0.41 (0.40); anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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, Liturgusa 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.

Type.

Holotype Male, pinned. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA.

Type locality.

Venezuela, Villa de Cura, AR. Venezuela, 1-VII-57, E. Doreste (Lat. 10.003890, Long. -67.476421).

Material examined.

Liturgusa cura sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Venezuela Villa de Cura, AR. Venezuela, 1-VII-57, E. Doreste 10.003890, -67.476421 ANSP 082
Female Allotype Venezuela MALUZ 004
Diagnosis.

The smallest Liturgusa species, Liturgusa cura is most similar to Liturgusa bororum and Liturgusa manausensis, but is located only in northern Venezuela, a unique distribution. Also, Liturgusa cura has rounded posterolateral margins of the metazone, distinct from Liturgusa manausensis, as well as a mostly pale ventral prothoracic femoral surface (male has a medial brown marking) that is distinguished from Liturgusa bororum.

Description.

Male. (Fig. 9A) N=1: Body length 18.74; forewing length 12.36; hindwing length 9.73; pronotum length 5.16; prozone length 1.57; pronotum width 2.19; pronotum narrow width 1.62; head width 4.42; head vertex to clypeus 1.78; frons width 1.51; frons height 0.52; prothoracic femur length 5.22; mesothoracic femur length 6.47; mesothoracic tibia length 4.83; mesothoracic tarsus length 4.56; metathoracic femur length 6.56; metathoracic tibia length 6.90; metathoracic tarsus length 6.38; pronotal elongation measure 0.30; pronotal shape measure 0.42; head shape measure 0.40; frons shape measure 0.34; anteroventral femoral spine count 15; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 9.

Liturgusa cura sp. n., dorsal habitus: A holotype male from Venezuela (ANSP 082) B allotype female from Venezuela (MALUZ 004).

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.

Female. (Fig. 9B) N=1: Body length 24.15; forewing length 14.24; hindwing length 10.75; pronotum length 6.84; prozone length 2.00; pronotum width 2.76; pronotum narrow width 2.09; head width 5.72; head vertex to clypeus 2.36; frons width 2.16; frons height 0.89; prothoracic femur length 6.32; mesothoracic femur length 7.68; mesothoracic tibia length 6.18; mesothoracic tarsus length 5.23; metathoracic femur length 7.57; metathoracic tibia length 8.61; pronotal elongation measure 0.29; pronotal shape measure 0.40; head shape measure 0.41; frons shape measure 0.41; anteroventral femoral spine count 15; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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, Liturgusa cura is named from the Villa de Cura in northern Venezuela where the species was collected.

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA.

Type locality.

Panama CZ, Madden Res., May 11‚ 72, R&E Froeschner (Lat. 9.119892, Long. -79.619867).

Material examined.

Liturgusa fossetti sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Panama CZ, Madden Res., May 11 ‚72, R&E Froeschner, USNM 013 9.119892, -79.619867 GSMC003836
Female Allotype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003425
Male Paratype Panama Barro Colo Isld., Canal Zone. I-7-1929, Collector C.H. Curran 9.164966, -79.837098 AMNH 040
Female Paratype Costa Rica ANSP 100
Female Paratype Panama Changuinola, 1-III-1917, C.B. Williams, on cocoa 9.477669, -82.473596 BMNH 087
Male Paratype Panama Gatun C.Z., March 1930, TO Zachokke, Tres Rios Plantation 9.270651, -79.909661 CAS 020
Male, Female Paratype Costa Rica Puntarenas Prov. Osa pen., nr. Carate, 8.43458°N, 83.38195°W, 10m, 01.VI.2010, gen. coll. 2010 ent. Class colr. KBM01061004 8.434580, -83.381956 GSMC003075-76
2 Males, 2 Females Paratype Costa Rica Puntarenas Prov. Osa pen., nr. Metapalo, 8.40765°N, 83.2820166°W, 14m, 29.V.2010, hand coll., 2010 ent. class colr. KBM02061002 8.407650, -83.282017 GSMC003078, GSMC003080-82
Male Paratype Panama Darien, F. Geay, 1899 8.092070, -77.722150 MNHN 069
Diagnosis.

A medium size species with a moderately elongate pronotum, Liturgusa fossetti is most similar to Liturgusa maya, which is also distributed in Central America. However, Liturgusa fossetti can easily be distinguished from Liturgusa maya and other Central American species by the yellow coloration on the hindwings.

Description.

Male. (Figs 2A, 10A) N=3: Body length 21.36–22.18 (21.76); forewing length 13.60–14.55 (14.21); hindwing length 10.26–11.53 (11.04); pronotum length 6.37–6.59 (6.49); prozone length 1.92–2.08 (1.98); pronotum width 2.47–2.54 (2.50); pronotum narrow width 1.63–1.84 (1.76); head width 4.96–5.13 (5.05); head vertex to clypeus 1.87–2.06 (1.95); frons width 1.73–1.78 (1.76); frons height 0.66–0.76 (0.69); prothoracic femur length 6.54–6.75 (6.62); mesothoracic femur length 7.92–8.27 (8.04); mesothoracic tibia length 5.94–6.28 (6.08); mesothoracic tarsus length 5.38–5.71 (5.54); metathoracic femur length 8.01–8.27 (8.14); metathoracic tibia length 7.96–8.83 (8.39); metathoracic tarsus length 7.43–8.20 (7.82); pronotal elongation measure 0.30–0.32 (0.30); pronotal shape measure 0.38–0.39 (0.39); head shape measure 0.37–0.40 (0.39); frons shape measure 0.38–0.43 (0.40); anteroventral femoral spine count 13–15 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 10.

Liturgusa fossetti sp. n., dorsal habitus: A holotype male from Panama (CLEV GSMC003836) B allotype female from Nicaragua (CLEV GSMC003425).

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.

Genital Complex (Fig. 51F.1): The main body of ventral left sclerite (L4A) slightly elongate with margins that taper rapidly to a medially pointed terminus, the left side highly sclerotized, the right membranous; 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) broad and heavily sclerotized with rapidly tapering margins terminating with a dull point, the concave margin strongly defined; the apical process (paa) thick and with a pronounced bulge at the base, curved and terminating with an evenly 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, broadened proximally with a few defined grooves; the ventral process (pva) c-shaped and broad, both ends rounded and blunt.

Female. (Figs 2B, 10B) N=3: Body length 30.16–32.36 (31.60); forewing length 18.38–20.15 (19.06); hindwing length 14.25–15.27 (14.76); pronotum length 9.22–9.53 (9.34); prozone length 2.93–2.95 (2.94); pronotum width 3.52–3.70 (3.60); pronotum narrow width 2.47–2.72 (2.60); head width 6.69–7.06 (6.84); head vertex to clypeus 2.75–3.00 (2.84); frons width 2.39–2.67 (2.54); frons height 0.91–1.10 (0.99); prothoracic femur length 8.85–9.75 (9.36); mesothoracic femur length 9.78–9.97 (9.88); mesothoracic tibia length 7.56–8.11 (7.83); mesothoracic tarsus length 6.94–7.11 (7.02); metathoracic femur length 9.55–9.92 (9.75); metathoracic tibia length 10.74–10.96 (10.87); metathoracic tarsus length 9.27–9.41 (9.34); pronotal elongation measure 0.31–0.32 (0.32); pronotal shape measure 0.37–0.40 (0.39); head shape measure 0.41–0.42 (0.41); frons shape measure 0.37–0.41 (0.39); anteroventral femoral spine count 15; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Etymology.

A noun in the genitive case, Liturgusa fossetti is named in honor of James Stephen Fossett for his inspirational dedication to adventure and exploration

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA.

Type locality.

Bolivia: Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson (Lat. -17.671350, Long. -63.457517).

Material examined.

Liturgusa kirtlandi sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000281
Female Allotype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000274
7 Males, 6 Females Paratype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000252-53, GSMC000267, GSMC000271, GSMC000275-76, GSMC000279-80, GSMC000282-84, GSMC000306-07
2 Females Paratype Bolivia Dpto. Santa Cruz, nr. Buena Vista, 17°39.100'S, 063°35.136'W, 390 m, 14.XI.2009, at MV.UV lights in cut trees, Coll: G.J. Svenson -17.651667, -63.585600 GSMC000272, GSMC000328
Diagnosis.

Most similar to Liturgusa maya and Liturgusa trinidadensis, but easily diagnosed by the elongate, thin apical process (paa) with an angled blunt tip rather than an evenly rounded terminus as is seen in the other two species. See Liturgusa trinidadensis for a list of external features that distinguish the species from Liturgusa kirtlandi. The main difference between Liturgusa maya and Liturgusa kirtlandi is the presence of tubercles on the pronotum in Liturgusa kirtlandi and not in Liturgusa maya. Finally, Liturgusa trinidadensis has numerous tubercles in the posterolateral corners of the metazone while Liturgusa kirtlandi does not.

Description.

Male. (Fig. 11A) N=4: Body length 25.82–26.83 (26.2); forewing length 17.37–17.91 (17.64); hindwing length 13.49–14.19 (13.86); pronotum length 6.95–7.35 (7.18); prozone length 2.06–2.26 (2.18); pronotum width 2.63–2.78 (2.70); pronotum narrow width 1.96–2.02 (1.99); head width 5.35–5.62 (5.47); head vertex to clypeus 2.14–2.21 (2.18); frons width 1.91–2.07 (1.98); frons height 0.61–0.77 (0.67); prothoracic femur length 6.76–7.04 (6.90); mesothoracic femur length 8.46–8.88 (8.71); mesothoracic tibia length 6.73–7.09 (6.94); mesothoracic tarsus length 5.80–6.10 (5.94); metathoracic femur length 8.54–9.12 (8.83); metathoracic tibia length 10.01–10.50 (10.33); metathoracic tarsus length 8.53–8.84 (8.70); pronotal elongation measure 0.30–0.31 (0.30); pronotal shape measure 0.36–0.39 (0.38); head shape measure 0.39–0.40 (0.40); frons shape measure 0.31–0.39 (0.34); anteroventral femoral spine count 13–14 (14); anteroventral tibial spine count 9; posteroventral tibial spine count 7.

Figure 11.

Liturgusa kirtlandi sp. n., dorsal habitus: A holotype male from Santa Cruz, Bolivia (CLEV GSMC000281) B allotype female from Santa Cruz, Bolivia (CLEV GSMC000274).

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, 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) 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.

Female. (Figs 1B, 11B) N=5: Body length 30.11–32.79 (30.94); forewing length 20.14–21.41 (20.55); hindwing length 15.17–16.86 (15.84); pronotum length 8.16–8.70 (8.43); prozone length 2.44–2.66 (2.57); pronotum width 3.12–3.31 (3.23); pronotum narrow width 2.36–2.41 (2.39); head width 6.31–6.76 (6.60); head vertex to clypeus 2.78–2.97 (2.85); frons width 2.46–2.71 (2.56); frons height 0.78–0.93 (0.88); prothoracic femur length 8.13–8.53 (8.25); mesothoracic femur length 9.27–9.84 (9.50); mesothoracic tibia length 7.70–8.16 (7.85); mesothoracic tarsus length 6.23–6.69 (6.43); metathoracic femur length 9.39–9.87 (9.58); metathoracic tibia length 11.41–11.75 (11.57); metathoracic tarsus length 9.16–9.79 (9.59); pronotal elongation measure 0.30–0.31 (0.30); pronotal shape measure 0.37–0.40 (0.38); head shape measure 0.42–0.45 (0.43); frons shape measure 0.31–0.36 (0.34); anteroventral femoral spine count 13–14 (13); anteroventral tibial spine count 9–10 (9); posteroventral tibial spine count 7.

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, Liturgusa 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.

Type.

Holotype Male, pinned. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Type locality.

Brazil: Amazonas, 18.1 km e Campinas field sta. Km 60 n Manaus, 22 Feb 1979, 02°30'S, 060°15'W, Terra firme forest canopy fogged with Pyrethrum Sample # 12, Montgomery, Erwin, Schimmel, Krischik, Date, Bacon colls (Lat. -2.500000, Long. -60.250000).

Material examined.

Liturgusa manausensis sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Brazil Amazonas, 18.1 km e Campinas field sta. Km 60 n Manaus, 22 Feb 1979, 02°30'S, 060°15'W, Terra firme forest canopy fogged with Pyrethrum Sample # 12, Montgomery, Erwin, Schimmel, Krischik, Date, Bacon colls. -2.500000, -60.250000 USNM 001; USNM ENT 00873997
Diagnosis.

One of the smaller species, Liturgusa 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.

Description.

Male. (Fig. 12A) N=1: Body length 19.40; forewing length 12.93; hindwing length 9.66; pronotum length 5.68; prozone length 1.62; pronotum width 2.07; pronotum narrow width 1.47; head width 4.66; frons width 1.59; frons height 0.54; prothoracic femur length 5.54; mesothoracic femur length 7.27; mesothoracic tibia length 5.72; mesothoracic tarsus length 4.97; metathoracic femur length 7.52; metathoracic tibia length 7.95; metathoracic tarsus length 7.83; pronotal elongation measure 0.28; pronotal shape measure 0.36; frons shape measure 0.34; anteroventral femoral spine count 15; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 12.

Liturgusa, dorsal habitus: A Liturgusa manausensis sp. n., holotype male from north of Manaus, Brazil (USNM 001) B Liturgusa maya Saussure & Zehntner, 1894, illustration of female from Peru by Julio Rivera.

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.

Liturgusa maya Saussure & Zehntner, 1894

http://species-id.net/wiki/Liturgusa_maya

Mantis annulipes (non Mantis annulipes Audinet Serville, 1838): von Charpentier 1843: 28–29, pl. 28 (partim).
Liturgousa annulipes: Saussure 1871: 102 (♂ only, partim).
Liturgousa cayennensis [Var.] maya: Saussure and Zehntner 1894: 160; Scudder 1901: 159, 407; Rehn 1903: 6; Marshall 1975: 318; Roy and Cuche 2008: 14, 21.
Liturgousa maya: Scudder 1901: 159, 419; Kirby 1904: 271; Hebard 1932: 211; Rehn 1935: 201, pl. 8, fig. 5; Hughes-Schrader 1950: 11–14, 38, 45, Table 1, Fig. 9; Hughes-Schrader 1951: 178, 180, 183–184, 186–187, Tables 1–2, Fig. 1; Hughes-Schrader 1953: 544–554; Henderson 1965: 215; Cerdá 1996: 76;
Liturgusa maya: Giglio-Tos 1927: 293; Beier 1935: 11; Jantsch 1991: 125; Terra 1995: 54; Jantsch 1999: 48; Maes and Roy 2000: 61; Lombardo and Agabiti 2001: 90, 96; Ehrmann 2002: 207; Agudelo 2004: 55, Table 3.1; Agudelo 2005: 3; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116, 142; Svenson and Whiting 2009: Appendix S1.
Liturgusa maja: Passerin d‘Entrèves 1981: 61.
= Liturgusa charpentieri: Giglio-Tos 1927: 294; Beier 1935: 11; Passerin d‘Entrèves 1981: 61; Terra 1995: 53; Salazar E. 1998: 105, Fig. 4; Jantsch 1999: 48; Salazar E. 1999: 10; Lombardo and Agabiti 2001: 90; Salazar E. 2002: 124; Ehrmann 2002: 207; Agudelo 2004: 55, Table 3.1; Agudelo 2005: 3; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116, 141. syn. n.
Type.

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

Type locality.

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

Material examined.

Liturgusa maya Saussure & Zehntner, 1894.

Sex Type Country Label Latitude Longitude Code
Male Lectotype Mexico Temax, N. Yucatan, Gaumer 21.143702, -88.942315 BMNH
Male Paralectotype Mexico Temax, N. Yucatan, Gaumer 21.143702, -88.942315 MHNG
Female Paralectotype Mexico Temax, N. Yucatan, Gaumer 21.143702, -88.942315 MHNG
Female Paralectotype Mexico Temax, N. Yucatan, Gaumer 21.143702, -88.942315 BMNH
Female Paralectotype Mexico Temax, N. Yucatan, Gaumer 21.143702, -88.942315 BMNH
Male nontype Colombia S.A. Felipe Ovalle, Q., Ac. 33501 AMNH 011
Male nontype Mexico Itzimna, Yucatan, IX-9-1964, Collectors J.C. & D. Pallister 21.025527, -89.682887 AMNH 020
Male nontype Venezuela Rancho Grande, nr. Maracay, Ven. 8-VII 1946 10.350000, -67.683330 AMNH 023
Male nontype Panama C.Z., Summit, XII - 1953, Collector N.L.H. Krauss 9.067496, -79.649077 AMNH 039
Male nontype Peru El Campamiente, Col Perene, Peru, 19 June 1920, Cornell Univ. Exp. -10.941437, -75.225954 ANSP 025
Male nontype Peru Quiroz, Rio Paucartambo -12.903073, -71.404748 ANSP 038
Male nontype Mexico San Rafael, Veracruz, Townsend 20.190622, -96.882611 ANSP 076
Male nontype Costa Rica ANSP 077
Female figure voucher (Rehn 1935) Costa Rica Surubres, near Santa Mater, (Poe.). 250m, I-1906, P. Biolley! 9.598454, -84.302680 ANSP 078
Female nontype Costa Rica Hamburg Farm, lower Rio Reventazon, C.R., VI.26.1936 (L. Navas), In banana field 10.250000, -83.450000 ANSP 079
Female nontype Costa Rica Pozo Azul 10.186150, -84.901512 ANSP 087
Female figure voucher (Rehn 1935) Nicaragua Mouth of Waspuk R. 14.631086, -84.438399 ANSP 088
Female nontype Panama Corozal, C.Z., XI.17.1913, Morgan Hebard ANSP 089
Male nontype Mexico Teapa, Tabasco, Feb. H.H.S. 17.556197, -92.943064 BMNH 009
Female nontype Honduras Cortes Provence, San Pedro Sula, foothills, ca. 800 ft., Stop 82-16, 5-13 May 1982, D.B. Weissman 15.475779, -88.054841 CAS 002
Male nontype Peru Monson Valley, Tingo Maria, XII-2-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 009
Female nontype Peru Monson Valley, Tingo Maria, XII-2-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 013
Female nontype Panama Ft. Clayton, C.Z., XI-44, K.E. Frick Collector 9.005408, -79.574522 CAS 022
Female nontype Colombia Buga - Valle, Bosque del Vinculo, 997m, 3°50'13.0"N, 76°18'00.0"W, 4-I-2006, Coll: T. Kondo 3.836944, -76.300000 GSMC000277
Female nontype Colombia Cali - Valle, Chorro de Platu, 28-XII-2005, Coll: T. Kondo GSMC000327
Female nontype Costa Rica Puntarenas Prov. Osa pen., nr. Carate; gen. coll. 2010 ent. Class colr. KBM 01061004 8.434580, -83.381956 GSMC003014
Male nontype Guatemala Izabal, Rio Dulce, Hotel Tijax. Collected along trails through old secondary growth tropical forest under rocks, logs & bark. N 15°40'12.2"W, 89°00'27.0", Elev: 49m, July 8, 2006. Coll: J. Huff, C. Viquez & D. Ortiz. Site 8. 15.670056, -89.007500 GSMC003070
Female nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003426
Female nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003427
Female nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003428
Male nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003429
Male nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003430
Male nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003431
Female nontype Nicaragua Granada, Reserva Silvestre Privada Domatila, 11.709000°N, 85.953500°W, 68m, 6-9 November 2010, Coll: Gavin J Svenson 11.709000, -85.953500 GSMC003432
Male nontype Nicaragua Rio San Juan Dpto., Refugio Bartola; KBM 18051201 10.972540, -84.338990 GSMC003597
Female nontype Venezuela Dto Ptumba, Edo Zulia, Fecha 5-8-90, Collec Luis, U. 8.936828, -72.163996 MALUZ 001
Female nontype Venezuela Dto Trujillo, Edo Trujillo, Fecha 20-3-89, Collec Eladio, K. 9.373003, -70.429807 MALUZ 002
Female nontype Peru Cusco, Quillabamba, 3.VII.1996, W. Catalan leg., UA 684-1996 -12.861863, -72.699170 MEKRB 002
Male nontype Peru Junin, Satipo. 16.V.1999, J. Laos leg. UA 1278-1999 -11.238653, -74.627567 MEKRB 003
Female nontype Peru Timbes, Papayal, 2000, P. Castillo leg., UA 021-2000 -4.100794, -80.677040 MEKRB 006
Female nontype Ecuador Cotopaxi, La Mana (Bosque secord), 7.VIII.2003, Rivera y Montgemuy, UA 103-2003 -0.934058, -79.218475 MEKRB 007
Female nontype Ecuador Cotopaxi, La Mana (Bosque secord), 7.VIII.2003, Rivera y Montgemuy, UA 103-2003 -0.934058, -79.218475 MEKRB 008
Female nontype Ecuador Cotopaxi, La Mana (Bosque secord), 7.VIII.2003, Rivera y Montgemuy, UA 103-2003 -0.934058, -79.218475 MEKRB 009
Female nontype Ecuador Cotopaxi, La Mana (Bosque secord), 7.VIII.2003, Rivera y Montgemuy, UA 103-2003 -0.934058, -79.218475 MEKRB 011
Female nontype Colombia Dept. Amazonas, Rio Igara Parana, 30 km aval La Chorrera, VI - VII 1974, M. Descamps rec. -1.197385, -72.937475 MNHN 016
Female nontype Colombia Dept. Amazonas, Rio Igara Parana, 30 km aval La Chorrera, VI - VII 1974, M. Descamps rec. -1.197385, -72.937475 MNHN 018
Female nontype Colombia Dept. Narino Entre Espriella Tumaco et Camp Experimental de I.F.A., 300m, 19-XI-1968, M. Descamps rec. 1.788077, -78.778943 MNHN 028
Male nontype Nicaragua Zelaya El Recreo Foret, X-1984, C. Amedegnato, S. Poulain Rec. 12.178332, -84.344478 MNHN 078
Female nontype Honduras Atlantida, Massif de Pico Bonita, 250m, VI/1995, T. Porion G. lachaume 15.692101, -86.853643 MNHN 085
Female nontype Mexico Col. of J. Amith 80439 (2011-06-17), 16°48'57"N, 98°41'25"W, 600 m, Yoloxochitl, Mpio, S.L. Acatlan, Gro. 16.815833, -98.690278 JAC
Female nontype Mexico Col. of J. Amith 80439 (2011-06-17), 16°48'57"N, 98°41'25"W, 600 m, Yoloxochitl, Mpio, S.L. Acatlan, Gro. 16.815833, -98.690278 JAC
Female nontype Peru Madre de Dios, Tambopata Nat‘l Reserve, -12.8368° -69.2932°, 250m, 2005, Coll: G.J. Svenson -12.836800, -69.293200 GSMC003005
Female nontype Peru Madre de Dios, Tambopata Nat‘l Reserve, -12.8368° -69.2932°, 250m, 22-29.V.2011, Colr. UNM collections class (K.B. Miller) -12.836800, -69.293200 CLEV
Female nontype Ecuador Napo: Tiputini, Biodiversity Stn., vic. Yasuni Natl. Pk. 14-18 Feb 1999, 0°38'S, 76°10'W. DC Darling, ROM 991050 -0.633333, -76.166667 ROM 002
Female nontype Guatemala SDEI 004
Male nontype Guatemala Polochic R, 22.3, Guat., Barber & Schwarz Coll 15.336297, -89.732437 USNM 014; USNM ENT 00873004
Female nontype Peru Tingo Maria, IV, V - 1952, P. Araoz -9.314153, -76.006745 USNM 035; USNM ENT 00873012
Female nontype Guatemala 25, L. Thiel, S. Sebastian, Retalhuleu 14.567871, -91.649246 USNM 037; USNM ENT 00873013
Female nontype Panama Cabima, May 17. 11, August Busck 9.129759, -79.527743 USNM 038; USNM ENT 00873014
Male nontype Panama Pedro Diaz Farm, Las Sabanas, Nov. 1, 1918. G-349, Dietz and Zetek. 8.573002, -80.681300 USNM 053; USNM ENT 00873010
Female nontype Costa Rica Quepos, C.R., III-17-1958, M.J. Stelzer, No. 58-17, on cacao 9.429236, -84.163056 USNM 067; USNM ENT 00873011
Female nontype Mexico Tapachula, Chiapas 14.908892, -92.241002 ZMHB 002
Female nontype Costa Rica Farm Hamburg, am Reventazon, 2.II.1932, F. Nevermann leg., Eing. Nr. 74. 1932 10.250000, -83.450000 ZMUH 018
Female nontype Costa Rica Farm Hamburg, am Reventazon, 2.Juli.1934. F. Nevermann leg., Eing. Nr. 146. 1935 10.250000, -83.450000 ZMUH 019
Female nontype Peru Madre de Dios; Rio Tambopata; Posada Amazonas; Canopy Tower; S12 48 16.6 W69 17 35.3; Sept. 2004; J.R. Cryan -12.804611, -69.293139 GSMC003062
Female nontype Ecuador 10.VI.67, rec. MAZ, cuenca, Ecuador, A.C. Allyn, Acc. 1969-20 -3.010204, -78.204794 FMNH 005
Female nontype Peru Tingo Maria, 670m, Hyanuce, Peru SA, Nov. 2006 -9.314153, -76.006745 FMNH 010
Taxonomic history.

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

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

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

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

Natural history.

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

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

Diagnosis.

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

Description.

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

Figure 13.

Liturgusa maya Saussure & Zehntner, 1894, dorsal habitus: A male from Nicaragua (CLEV GSMC003431) B female from Costa Rica (ZMUH 018).

Figure 14.

Liturgusa maya Saussure & Zehntner, 1894, dorsal habitus and labels: A lectotype male (BMNH) B labels of lectotype male C paralectotype female (BMNH) D labels of paralectotype female.

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

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

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

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

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

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

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

Redescription.

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

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

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

Prothoracic Legs: As described for males.

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

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

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

Liturgusa stiewei sp. n.

http://zoobank.org/0EB1C2D3-98D5-4315-989F-7DFB5D319504

http://species-id.net/wiki/Liturgusa_stiewei

Liturgousa mesopoda: Hebard 1919b: 134.
Type.

Holotype Female, pinned. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA.

Type locality.

Jimenez, W. Colombia, elev. 1600ft. III 1907, M.G. Palmer (Lat. 2.675931, Long. -77.148414)

Material examined.

Liturgusa stiewei sp. n.

Sex Type Country Label Latitude Longitude Code
Female Holotype Colombia Jimenez, W. Colombia, elev. 1600ft. III 1907, M.G. Palmer 2.675931, -77.148414 ANSP 017
Male Allotype Colombia Santander Province 6.641759, -73.653934 MSMC
Female Paratype Colombia ANTIOQUIA: Maceo, to 1600 masl, ♀, 16-XII-2003, E. Henao leg (CJASE, Manizales) 6.550551, -74.778359 CJASE
Diagnosis.

One of the largest species of Liturgusa, comparable to Liturgusa nubeculosa, Liturgusa stiewei is more comparable to many large species of Hagiomantis. Can be distinguished from Liturgusa nubeculosa by its shorter, more broad pronotum and black, more opaque hindwings.

Description.

Male from copal (restricted access to some characters). (Fig. 15A) N=1: Body length 30.70; forewing length 19.10; pronotum length 9.18; prozone length 3.10; pronotum width 3.37; pronotum narrow width 2.33; head width 5.90; prothoracic femur length 7.75; mesothoracic femur length 12.14; mesothoracic tibia length 10.07; mesothoracic tarsus length 7.30; metathoracic femur length 12.96; metathoracic tibia length 13.20; metathoracic tarsus length 13.12; pronotal elongation measure 0.34; pronotal shape measure 0.37; anteroventral femoral spine count 14; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 15.

Liturgusa stiewei sp. n., dorsal habitus: A allotype male in copal (MSMC) B holotype female from Colombia (ANSP 017).

Head: Transverse, the juxta-ocular protuberances small, but pronounced, the apex in the middle; the vertex near straight, even with the dorsal margin of the eyes. Frontal suture with a slight medial carina forming a continuous arc. Ocelli small. Frons, clypeus and labrum not visible for description. Palpi are pale.

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.

Genital Complex: Only male in copal.

Female. (Fig. 15B) N=2: Body length 41.94–42.00 (41.97); forewing length 26.10–26.70 (26.40); hindwing length 21.20; pronotum length 11.14–11.15 (11.14); prozone length 2.60–3.16 (2.88); pronotum width 4.16; pronotum narrow width 2.97; head width 7.24; head vertex to clypeus 3.16; frons width 2.81; frons height 1.06; prothoracic femur length 11.86–12.10 (11.98); mesothoracic femur length 14.59; mesothoracic tibia length 9.53; metathoracic femur length 13.70–14.00 (13.85); metathoracic tibia length 15.67; metathoracic tarsus length 13.20–13.42 (13.31); pronotal elongation measure 0.23–0.28 (0.26); pronotal shape measure 0.37; head shape measure 0.44; frons shape measure 0.38; anteroventral femoral spine count 16; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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 dorso-lateral 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.

Etymology.

A noun in the genitive case, Liturgusa stiewei is named for Martin Stiewe in honor of his contributions to Mantodea systematics and his collaboration in discovering this new species.

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA

Type locality.

Bolivia: La Paz Dept., Ituralde Prov., San Miguel, 14°30.602'S, 67°29.555'W, 24-30 Sept 2007, Coll: K.B. Miller et al. (Lat. -14.510033, Long. -67.492583)

Material examined.

Liturgusa tessae sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Bolivia La Paz Dept., Ituralde Prov., San Miguel, 14°30.602'S, 67°29.555'W, 24-30 Sept 2007, Coll: K.B. Miller et al. -14.510033, -67.492583 GSMC000263
Female Allotype Peru Smithsonian Earthwatch Sample, Location 8j, Nickle, D. GSMC000265
Female Paratype Brazil Santarem, July. 1919, S.M. Klages. Acct. 6324 -2.465914, -54.701313 ANSP 111
Female Paratype Brazil Santarem, July. 1919, S.M. Klages. Acct. 6324 -2.465914, -54.701313 ANSP 108
Male Paratype Brazil Santarem, July. 1919, S.M. Klages. Acct. 6324 -2.465914, -54.701313 ANSP 109
Male Paratype Brazil Santarem, July. 1919, S.M. Klages. Acct. 6324 -2.465914, -54.701313 ANSP 110
Female Paratype Brazil Santarem, July. 1919, S.M. Klages. Acct. 6324 -2.465914, -54.701313 ANSP 112
Female Paratype Brazil Santarem, Oct. 1919, Carn. Mus. Acc. 6543 -2.465914, -54.701313 ANSP 113
Male Paratype Brazil 50, 2 BMNH 002
Male Paratype Brazil 52, 96 BMNH 005
Female Paratype Peru Rio Tambopata, Explorer‘s Inn-Rio Tower, 12°50.208'S, 069°17.603'W, 10-XII-2003, Svenson -12.836800, -69.293383 GSMC000261
Male Paratype Bolivia La Paz Dept., Ituralde Prov., San Miguel, 14°30.602'S, 67°29.555'W, 24-30 Sept 2007, Coll: K.B. Miller et al. -14.510033, -67.492583 GSMC000268
Female Paratype Brazil Amazonas Reserva Biologica do Cuieiras, 50 km. N. Manaus, 15-IV-AU 15-V-1981, M. Descamps -2.599556, -60.210631 MNHN 020
Male Paratype Brazil Para, Ile de Mosqueiro, 11-9-83, male + female accouples sur trone -1.157643, -48.465362 MNHN 086
Female Paratype Brazil Para, Ile de Mosqueiro, 11-9-83, male + female accouples sur trone -1.157643, -48.465362 MNHN 087
Male Paratype Brazil Para: Rio Xingu, Camp (52°22'W, 3°39'S), ca 60 km S. Altamira, 8-12 Oct 1986, P. Spangler & O. Flint -3.650000, -52.366667 USNM 007; USNM ENT 00873005
Male Paratype Brazil Para: Rio Xingu, Camp (52°22'W, 3°39'S), ca 60 km S. Altamira, 8-12 Oct 1986, P. Spangler & O. Flint -3.650000, -52.366667 USNM 008; USNM ENT 00873006
Male Paratype Brazil Para. ZMHB 004
Male Paratype Peru Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 07 May 89, 05/02/051 -12.833333, -69.283333 USNM 069; USNM ENT 00873048
Female Paratype Peru Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonado, 290 m, 12°50'S, 69°17'W; Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al., colls. 08 Nov 83 May 89, 04/01/072 -12.833333, -69.283333 USNM 070; USNM ENT 00873049
Male Paratype Bolivia La Paz Dept., Ituralde Prov., San Miguel, 14°30.602'S, 67°29.555'W, 24-30 Sept 2007, Coll: K.B. Miller et al. -14.510033, -67.492583 GSMC000317
Diagnosis.

Very similar to Liturgusa maya in size and shape, but with an even color patterning on the forewings, lacking highly contrasting regions. The hindwings for Liturgusa 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).

Description.

Male. (Fig. 16A) N=9: Body length 19.23–23.89 (22.12); forewing length 13.88–15.21 (14.61); hindwing length 10.95–12.13 (11.47); pronotum length 5.71–6.65 (6.13); prozone length 1.68–2.00 (1.82); pronotum width 2.36–2.62 (2.48); pronotum narrow width 1.64–1.92 (1.76); head width 4.70–5.51 (5.03); head vertex to clypeus 1.90–2.16 (1.97); frons width 1.64–1.95 (1.77); frons height 0.61–0.69 (0.65); prothoracic femur length 5.73–6.64 (6.12); mesothoracic femur length 6.92–8.47 (7.67); mesothoracic tibia length 5.54–6.65 (5.97); mesothoracic tarsus length 4.84–5.70 (5.17); metathoracic femur length 7.23–8.63 (7.80); metathoracic tibia length 7.78–9.23 (8.27); metathoracic tarsus length 7.25–8.35 (7.61); pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.39–0.43 (0.41); head shape measure 0.37–0.40 (0.39); frons shape measure 0.35–0.42 (0.37); anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 9–10 (10); posteroventral tibial spine count 7.

Figure 16.

Liturgusa tessae sp. n., dorsal habitus: A holotype male from San Miguel, Bolivia (CLEV GSMC000263) B allotype female from Tambopata, Peru (CLEV GSMC000265).

Head (Fig. 42C): Transverse, the juxta-ocular protuberances small, the apex in the lateral half; the vertex slightly concave, slightly above the dorsal margin of the eyes. 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.

Female. (Fig. 16B) N=7: Body length 23.65–29.63 (26.24); forewing length 14.71–18.43 (16.76); hindwing length 12.34–13.05 (12.7); pronotum length 6.77–7.86 (7.10); prozone length 1.99–2.27 (2.12); pronotum width 2.74–3.16 (2.93); pronotum narrow width 1.99–2.37 (2.14); head width 5.65–6.34 (5.98); head vertex to clypeus 2.35–2.64 (2.51); frons width 2.20–2.55 (2.38); frons height 0.74–0.98 (0.84); prothoracic femur length 6.68–7.83 (7.08); mesothoracic femur length 7.72–9.45 (8.33); mesothoracic tibia length 5.87–7.49 (6.57); mesothoracic tarsus length 5.23–6.39 (5.68); metathoracic femur length 7.80–9.66 (8.54); metathoracic tibia length 8.46–10.72 (9.46); metathoracic tarsus length 7.89–9.15 (8.34); pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.39–0.43 (0.41); head shape measure 0.41–0.44 (0.42); frons shape measure 0.33–0.39 (0.35); anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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: Moderately widened.

Etymology.

A noun in the genitive case, Liturgusa tessae is named for my daughter Tessa Eliza Svenson.

Liturgusa trinidadensis sp. n.

http://zoobank.org/56891F11-5672-4A7F-9D93-B6C0F4CCD515

http://species-id.net/wiki/Liturgusa_trinidadensis

Liturgousa maya (partim): Rehn 1935: 202.
Type.

Holotype Male, pinned. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA.

Type locality.

Trinidad, 17 I‚ 52, F. Schrader, 709 (Lat. 10.240326, Long. -61.217020).

Material examined.

Liturgusa trinidadensis sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Trinidad 17 I ‚52, F. Schrader, 709 10.240326, -61.217020 ANSP 035
Female Allotype Trinidad Caparo, VIII 1913, S.M. Klages 10.449908, -61.333566 ANSP 099
Male Paratype Trinidad W.I., Trinidad: Arima Vall. A.W.N.C. 7/15/78, R.A. Mendez 10.661851, -61.289723 AMNH 029
Female Paratype Trinidad 10 I ‚52, F. Schrader, 697 10.240326, -61.217020 ANSP 001
Male Paratype Trinidad 18 I ‚52, F. Schrader, 715 10.240326, -61.217020 ANSP 002
Male Paratype Trinidad 17 I ‚52, F. Schrader, 711 10.240326, -61.217020 ANSP 003
Male Paratype Trinidad 18 I ‚52, F. Schrader, 723 10.240326, -61.217020 ANSP 004
Male Paratype Trinidad 11 I ‚52, F. Schrader, 781 10.240326, -61.217020 ANSP 005
Female Paratype Trinidad 18 I ‚52, F. Schrader, 725 10.240326, -61.217020 ANSP 006
Female Paratype Trinidad 18 I ‚52, F. Schrader, 724 10.240326, -61.217020 ANSP 007
Male Paratype Trinidad 17 I ‚52, F. Schrader, 710 10.240326, -61.217020 ANSP 008
Male Paratype Trinidad 17 I ‚52, F. Schrader, 712 10.240326, -61.217020 ANSP 009
Male Paratype Trinidad 15 I ‚52, F. Schrader, 699 10.240326, -61.217020 ANSP 014
Male Paratype Trinidad 18 I ‚52, F. Schrader, 722 10.240326, -61.217020 ANSP 015
Male Paratype Trinidad 17 I ‚52, F. Schrader, 713 10.240326, -61.217020 ANSP 022
Female Paratype Trinidad 17 I ‚52, F. Schrader, 714 10.240326, -61.217020 ANSP 023
Female Paratype Trinidad Arima Valley, B.W.I. 6-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 024
Male Paratype Trinidad 18 I ‚52, F. Schrader, 721 10.240326, -61.217020 ANSP 026
Female Paratype Trinidad Arima Valley, B.W.I. 7-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 027
Female Paratype Trinidad Arima Valley, B.W.I. 10-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 028
Male Paratype Trinidad Arima Valley, B.W.I. 10-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 029
Male Paratype Trinidad Arima Valley, B.W.I. 7-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 030
Female Paratype Trinidad 19 I ‚52, F. Schrader, 717 10.240326, -61.217020 ANSP 032
Male Paratype Trinidad Nov. 1940, St. Augustine, BWI, H.S. Darling 10.659048, -61.398679 ANSP 034
Male Paratype Trinidad Arima Valley, B.W.I. 7-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 036
Male Paratype Trinidad Arima Valley, B.W.I. 10-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 ANSP 037
Male Paratype Trinidad Caparo, VI 1913, S.M. Klages 10.449908, -61.333566 ANSP 083
Male Paratype Trinidad Caparo, VI 1913, S.M. Klages 10.449908, -61.333566 ANSP 084
Male Paratype Trinidad Caparo, VI 1913, S.M. Klages 10.449908, -61.333566 ANSP 085
Female Paratype Trinidad Caparo, VIII 1913, S.M. Klages 10.449908, -61.333566 ANSP 086
Female Paratype Trinidad Caparo, VIII 1913, S.M. Klages 10.449908, -61.333566 ANSP 095
Female Paratype Trinidad Caparo, VIII 1913, S.M. Klages 10.449908, -61.333566 ANSP 096
Female Paratype Trinidad Caparo, VI 1913, S.M. Klages 10.449908, -61.333566 ANSP 097
Male Paratype Trinidad Moruga Area, ii-iii.1986, G.B. Popov 10.089796, -61.279058 BMNH 081
Female Paratype Trinidad Moruga Area, ii-iii.1986, G.B. Popov 10.089796, -61.279058 BMNH 082
Male Paratype Trinidad Moruga Area, ii-iii.1986, G.B. Popov 10.089796, -61.279058 BMNH 083
Female Paratype Trinidad Moruga Area, ii-iii.1986, G.B. Popov 10.089796, -61.279058 BMNH 084
Male Paratype Trinidad Arima Valley, B.W.I. 20-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 USNM 017; USNM ENT 00873007
Female Paratype Trinidad Arima Valley, B.W.I. 8-II-1952, Tropical Research Station, New York Zool Society 10.661851, -61.289723 USNM 018; USNM ENT 00873008
Male Paratype Trinidad Jun WI, Aug. Busck Collector USNM 019; USNM ENT 00873009
Male Paratype Trinidad Jun WI, Aug. Busck Collector USNM 020; USNM ENT 00873015
Male Paratype Trinidad Aug-22-1907, O.W. Barria, On Cacao, 252 USNM 052; USNM ENT 00873016
Diagnosis.

Most similar to Liturgusa maya and Liturgusa kirtlandi, exhibiting similar size, coloration and pronotum shape, Liturgusa 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 Liturgusa maya and Liturgusa kirtlandi have larger central ocelli. Finally, Liturgusa 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.

Description.

Male. (Fig. 17A) N=15: Body length 20.29–23.06 (21.63); forewing length 13.29–15.43 (14.09); hindwing length 11.61–11.90 (11.75); pronotum length 5.89–6.85 (6.19); prozone length 1.73–2.11 (1.87); pronotum width 2.30–2.80 (2.42); pronotum narrow width 1.53–2.06 (1.74); head width 4.70–5.43 (5.01); head vertex to clypeus 1.87–2.23 (2.00); frons width 1.71–1.97 (1.82); frons height 0.59–0.71 (0.65); prothoracic femur length 5.74–6.70 (6.10); mesothoracic femur length 6.87–7.89 (7.45); mesothoracic tibia length 5.49–6.39 (5.93); mesothoracic tarsus length 4.79–5.51 (5.22); metathoracic femur length 7.05–7.82 (7.48); metathoracic tibia length 7.65–9.08 (8.40); metathoracic tarsus length 6.95–7.91 (7.42); pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.38–0.41 (0.39); head shape measure 0.38–0.41 (0.40); frons shape measure 0.34–0.39 (0.36); anteroventral femoral spine count 14–16 (15); anteroventral tibial spine count 10–11 (10); posteroventral tibial spine count 7.

Figure 17.

Liturgusa trinidadensis sp. n., dorsal habitus: A holotype male from Trinidad (ANSP 035) B allotype female from Caparo, Trinidad (ANSP 099).

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.

Female. (Fig. 17B) N=11: Body length 22.56–31.05 (26.47); forewing length 13.35–17.48 (15.29); hindwing length 12.61; pronotum length 6.55–7.78 (7.30); prozone length 1.91–2.37 (2.22); pronotum width 2.63–3.09 (2.91); pronotum narrow width 2.04–2.25 (2.14); head width 5.37–6.41 (6.08); head vertex to clypeus 2.27–2.77 (2.59); frons width 2.12–2.52 (2.36); frons height 0.76–0.94 (0.86); prothoracic femur length 6.34–7.65 (7.15); mesothoracic femur length 7.20–8.73 (8.14); mesothoracic tibia length 5.78–7.42 (6.75); mesothoracic tarsus length 4.95–6.42 (5.76); metathoracic femur length 7.16–8.80 (8.11); metathoracic tibia length 8.13–10.95 (9.68); metathoracic tarsus length 7.30–8.75 (8.02); pronotal elongation measure 0.29–0.32 (0.30); pronotal shape measure 0.38–0.41 (0.40); head shape measure 0.40–0.45 (0.43); frons shape measure 0.34–0.39 (0.37); anteroventral femoral spine count 14–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Type.

Holotype Male, pinned. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Type locality.

Guatemala, Alta V. Paz, Schwarz & Barber Coll, 2.4 Cacao, Trece Aguas. (Lat. 15.592321, Long. -90.146392).

Material examined.

Liturgusa zoae sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Guatemala Alta V. Paz, Schwarz & Barber Coll, 2.4 Cacao, Trece Aguas. 15.592321, -90.146392 USNM 062; USNM ENT 00873990
Female Allotype Guatemala Alta V. Paz, Schwarz & Barber Coll, 2.4 Cacao, Trece Aguas. 15.592321, -90.146392 USNM 045; USNM ENT 00873991
Male Paratype Honduras 192, Ac. 29596 AMNH 002
Female Paratype Panama Puerto Armuelles 8.274772, -82.865134 AMNH 028
Nymph nontype Honduras Atlantida, Massif Pico Bonito, env. El Pino 200 m, VII/1995 T. Porion A. Grange 15.716539, -86.826280 MNHN 200
Female Paratype Belize Toledo Dist., Blue Creek Village, 22 June 1981, W.E. Steiner 16.195801, -89.043072 USNM 002; USNM ENT 00873017
Diagnosis.

A medium sized species, is distinct from all other Liturgusa. Could be considered most similar to Liturgusa guyanensis based on the pronounced pronotal shape modifications compared to all other Liturgusa, but Liturgusa 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.

Description.

Male. (Fig. 18A) N=3: Body length 23.28–25.87 (24.69); forewing length 15.09–17.00 (16.17); hindwing length 13.89; pronotum length 6.61–7.23 (6.85); prozone length 1.89–2.01 (1.96); pronotum width 2.47–3.00 (2.75); pronotum narrow width 1.89–1.98 (1.94); head width 4.95–5.52 (5.29); head vertex to clypeus 1.98–2.29 (2.18); frons width 1.66–2.00 (1.85); frons height 0.66–0.80 (0.75); prothoracic femur length 6.77–7.35 (7.01); mesothoracic femur length 8.02–9.37 (8.67); mesothoracic tibia length 6.37–7.08 (6.65); mesothoracic tarsus length 5.21–6.44 (5.82); metathoracic femur length 8.62–9.26 (8.94); metathoracic tibia length 8.84–9.81 (9.32); metathoracic tarsus length 9.47; pronotal elongation measure 0.28–0.29 (0.29); pronotal shape measure 0.37–0.41 (0.40); head shape measure 0.40–0.42 (0.41); frons shape measure 0.40–0.41 (0.40); anteroventral femoral spine count 14–15 (15); anteroventral tibial spine count 9; posteroventral tibial spine count 7.

Figure 18.

Liturgusa zoae sp. n., dorsal habitus: A holotype male from Guatemala (USNM 062) B allotype female from Guatemala (USNM 045).

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.

Female. (Fig. 18B) N=2: Body length 27.92; forewing length 17.64; hindwing length 13.94; pronotum length 7.65–8.30 (7.97); prozone length 2.22–2.59 (2.40); pronotum width 3.23–3.26 (3.25); pronotum narrow width 2.16–2.44 (2.30); head width 6.00–6.22 (6.11); head vertex to clypeus 2.44–2.65 (2.54); frons width 2.23–2.31 (2.27); frons height 0.90–0.97 (0.94); prothoracic femur length 7.98–8.17 (8.07); mesothoracic femur length 8.83–9.54 (9.18); mesothoracic tibia length 6.94–7.09 (7.02); mesothoracic tarsus length 6.38–6.68 (6.53); metathoracic femur length 9.15; metathoracic tibia length 9.93; metathoracic tarsus length 9.93; pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.39–0.43 (0.41); head shape measure 0.41–0.43 (0.42); frons shape measure 0.40–0.42 (0.41); anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 9–10 (9); posteroventral tibial spine count 7.

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, Liturgusa zoae is named for my daughter Zoey Kay Svenson.

Cursor Group A
Liturgusa cursor Rehn, 1950

http://species-id.net/wiki/Liturgusa_cursor

Liturgusa annulipes: Rehn 1935: 199, pl. 8, fig. 4.
Liturgousa cursor: Rehn 1950: 369–376, Figs 6–11; Hughes-Schrader 1950: 11–14, 27, 38, 45, Table 1, Fig. 11; Hughes-Schrader 1951: 178, 183–184, 186–187, Tables 1–2, Fig. 3; Hughes-Schrader 1953: 544–554; Henderson 1965: 215; Otte 1978: 76;
Liturgusa cursor: Terra 1995: 54; Jantsch 1999: 48; Ehrmann 2002: 207; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116.
Type.

Holotype Male. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA, Type no. 5761.

Type locality.

Panama, Barro Colorado Island, Gatun Lake, Canal Zone, December 31, 1948. (Dr. Franz Schrader, no. 630.) (Lat. 9.164966, Long. -79.837098).

Material examined.

Liturgusa cursor Rehn, 1950.

Sex Type Country Label Latitude Longitude Code
Male Holotype Panama Barro Colorado Island, Gatun Lake, Canal Zone, December 31, 1948. (Dr. Franz Schrader, no. 630.) 9.164966, -79.837098 ANSP
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 614 9.164966, -79.837098 ANSP 058
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 639 9.164966, -79.837098 ANSP 059
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 624 9.164966, -79.837098 ANSP 060
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 631 9.164966, -79.837098 ANSP 061
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 638 9.164966, -79.837098 ANSP 062
Male Paratype Panama Barro Colorado C.Z., 31-XII-‘48, F. Schrader, 653 9.164966, -79.837098 ANSP 063
Female Paratype Costa Rica Lower Rio Reventazon, Castilla Farm, VII. 23. 1936, C.W. Dodge 10.078445, -83.573754 ANSP 070
Male Paratype Costa Rica Turrialba, 8 VI, 48, F. Schrader, 515 9.989971, -83.763587 ANSP 071
Male Paratype Costa Rica ANSP 073
Female Paratype Costa Rica Ujarass de Terraba, C.R. IX.10.07, M.A.C. Jr. 9.831438, -83.832067 ANSP 074
Female nontype Panama Bugaba, 800-1, 500 ft., Champion., Godman-Salvin Coll. 1894-125. 8.490081, -82.620285 BMNH 008
Female nontype Panama Bocas del Toro Prov., Palo Seco NP, at ANAM station, 08°47.602'N, 082°11.330'W, 523m, 4.VI.2009, N. Lord, K. Miller, E. Nearns Colrs., General collecting 8.793367, -82.188833 GSMC000285
Female nontype Costa Rica Alajuela, Rio San Juan across from Bartola, 10.974153°N, 84.343499°W, 4 November, 2010, Coll: Gavin J. Svenson 10.974153, -84.343499 GSMC002952
25 Males, 18 Females nontype Nicaragua Rio San Juan, Refugio Bartola sur le Rio San Juan, 10.974309°N, 84.338318°W, 52 m, 1-5 November, 2010, Coll: Gavin J. Svenson 10.974309, -84.338318 GSMC003039, GSMC003043, GSMC003049, GSMC003055, GSMC003057, GSMC003060, GSMC003433-69
9 Males, 8 Females nontype Nicaragua Rio San Juan Dpto., Refugio Bartola; KBM 18051201 10.972540, -84.338990 GSMC003598, GSMC003602, GSMC003604, GSMC003607-08, GSMC003612-13, GSMC003621-22, GSMC003624-25, GSMC003627-28, GSMC003635-36, GSMC003646-47
Female nontype Nicaragua Rio San Juan, Cerro El Gigante, IX-98, col. I. Coronado 11.316590, -84.612565 MNHN 080
Female nontype Panama Barro Colo Is CZ, VI-18-37, S.W. Frost colr. 9.164966, -79.837098 USNM 049; USNM ENT 00873018
Female nontype Costa Rica Farm Hamburg, am Reventazon, 2.II.1932, F. Nevermann leg., Einz. Nr. 74. 1932 10.250000, -83.450000 ZMUH 003
Male nontype Costa Rica Farm Hamburg, am Reventazon, 2.II.1932, F. Nevermann leg., Einz. Nr. 74. 1932 10.250000, -83.450000 ZMUH 004
Female nontype Costa Rica Puntarenas Prov. Osa pen., nr. Matapalo; MV light coll. 2010 ent. Class colr. KBM 30051003 8.399580, -83.313750 GSMC003085
Female nontype Panama Barro C‘do I., I-20-26: 59, C.Z. 9.164966, -79.837098 FMNH 008
Male nontype Panama Barro Colorado I., Canal Zone, Panama, I-13-59, H.S. Dybas leg., at light 9.164966, -79.837098 FMNH 009
Taxonomic history.

Described in 1950 by James Rehn, the species was originally identified as Liturgusa annulipes by Rehn in his 1935 work on Orthoptera of Costa Rica. Rehn later recognized that the species was unique, recording specimens mainly from Barro Colorado in Panama, but including one specimen from Costa Rica. We found the species in southern Nicaragua as well, which is a new record for that country. 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 Liturgusa fossetti, Liturgusa maya, and Corticomantis atricoxata.

Diagnosis.

A small, slender species like Liturgusa dominica and Liturgusa milleri, but restricted in distribution to Central America. The forewings of males are colored asymmetrically while Liturgusa dominica and Liturgusa 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.

Redescription.

Male. (Fig. 19A) N=11: Body length 19.23–22.61 (20.94); forewing length 12.00–13.18 (12.57); hindwing length 9.34–9.84 (9.52); pronotum length 6.27–7.01 (6.66); prozone length 1.82–2.07 (1.99); pronotum width 1.99–2.17 (2.09); pronotum narrow width 1.32–1.54 (1.39); head width 4.37–4.74 (4.51); head vertex to clypeus 1.69–1.85 (1.76); frons width 1.39–1.56 (1.47); frons height 0.53–0.74 (0.60); prothoracic femur length 6.03–6.70 (6.33); mesothoracic femur length 8.41–9.37 (8.71); mesothoracic tibia length 6.42–7.28 (6.67); mesothoracic tarsus length 5.81–6.78 (6.31); metathoracic femur length 8.18–9.23 (8.56); metathoracic tibia length 8.68–9.83 (8.93); metathoracic tarsus length 9.72–11.46 (10.23); pronotal elongation measure 0.29–0.30 (0.30); pronotal shape measure 0.30–0.33 (0.32); head shape measure 0.38–0.40 (0.39); frons shape measure 0.38–0.47 (0.41); anteroventral femoral spine count 14–15 (15); anteroventral tibial spine count 9; posteroventral tibial spine count 7.

Figure 19.

Liturgusa cursor Rehn, 1950, dorsal habitus: A paratype male from Barro Colorado Island, Panama (ANSP 061) B paratype female from Barro Colorado Island, Panama (ANSP 070).

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 tooth-like; the ventral process (pva) c-shaped and smooth.

Redescription.

Female. (Figs 1A, 19B) N=11: Body length 24.15–29.52 (26.39); forewing length 13.83–16.18 (14.73); hindwing length 10.73–12.71 (11.46); pronotum length 7.37–8.32 (7.88); prozone length 2.20–2.49 (2.35); pronotum width 2.38–2.78 (2.58); pronotum narrow width 1.55–2.02 (1.74); head width 5.23–5.48 (5.35); head vertex to clypeus 2.19–2.38 (2.28); frons width 1.85–1.97 (1.90); frons height 0.71–0.86 (0.79); prothoracic femur length 7.19–7.97 (7.64); mesothoracic femur length 9.28–10.45 (9.84); mesothoracic tibia length 7.25–8.09 (7.73); mesothoracic tarsus length 6.73–7.43 (7.10); metathoracic femur length 9.25–10.53 (9.80); metathoracic tibia length 9.88–10.99 (10.33); metathoracic tarsus length 10.70–11.61 (11.11); pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.31–0.35 (0.33); head shape measure 0.41–0.46 (0.43); frons shape measure 0.38–0.44 (0.42); anteroventral femoral spine count 14–15 (15); anteroventral tibial spine count 9–10 (9); posteroventral tibial spine count 7.

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.

Type.

Holotype Male, pinned. National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Type locality.

Dominica, Grand Bay, IV-13-1964, O.S. Flint, Jr. (Lat. 15.239545, Long. -61.320099)

Material examined.

Liturgusa dominica sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 036: USNM ENT 00873995
Female Allotype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 034: USNM ENT 00873996
Female Paratype Dominica B.W.I., La Haut Estate, 18-I-1979, A.T. Finnamore Lyman Museum
Nymph Paratype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 039; USNM ENT 00873019
Female Paratype Dominica Fond Figues, III-16-1964, Dale F. Bray, Bredin-Archbold-Smithsonian Bio.Surv.Dominica USNM 040; USNM ENT 00873020
Male Paratype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 041; USNM ENT 00873021
Male Paratype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 042; USNM ENT 00873022
Male Paratype Dominica Grand Bay, IV-13-1964, O.S. Flint, Jr., Bredin-Archbold-Smithsonian Bio.Surv.Dominica 15.239545, -61.320099 USNM 043; USNM ENT 00873023
Nymph Paratype Dominica Clarke Hall, 22-31-X-1966, A.B. Gurney USNM 021; USNM ENT 00873024
Diagnosis.

This small species can be identified based on its slender size and its geographic restriction to the Caribbean island of Dominica. Much like Liturgusa 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 Liturgusa milleri and Liturgusa 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 Liturgusa dominica are restricted to French Guiana and Central America (Liturgusa milleri and Liturgusa cursor, respectively).

Description.

Male. (Fig. 20A) N=4: Body length 21.12–21.82 (21.38); forewing length 10.74–11.71 (11.29); hindwing length 8.12; pronotum length 6.91–7.28 (7.11); prozone length 2.02–2.11 (2.06); pronotum width 2.01–2.15 (2.09); pronotum narrow width 1.33–1.45 (1.40); head width 4.36–4.56 (4.47); head vertex to clypeus 1.55–1.67 (1.62); frons width 1.47–1.58 (1.53); frons height 0.56–0.60 (0.57); prothoracic femur length 6.02–6.47 (6.28); mesothoracic femur length 8.42–8.91 (8.66); mesothoracic tibia length 6.21–6.60 (6.44); mesothoracic tarsus length 5.79–6.19 (5.97); metathoracic femur length 8.58–9.05 (8.82); metathoracic tibia length 8.70–9.18 (8.93); metathoracic tarsus length 8.57–9.08 (8.80); pronotal elongation measure 0.29; pronotal shape measure 0.29–0.30 (0.29); head shape measure 0.36–0.38 (0.36); frons shape measure 0.36–0.40 (0.37); anteroventral femoral spine count 14–16 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7 (one male with 6 on left tibia).

Figure 20.

Liturgusa dominica sp. n., dorsal habitus: A holotype male from Dominica (USNM 036) B allotype female from Dominica (USNM 034).

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 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 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.

Female. (Fig. 20B) N=2: Body length 26.05–27.47 (26.76); forewing length 12.52–12.79 (12.65); hindwing length 9.29; pronotum length 8.67–9.04 (8.85); prozone length 2.50–2.66 (2.58); pronotum width 2.57–2.65 (2.61); pronotum narrow width 1.66–1.72 (1.69); head width 5.22–5.29 (5.26); head vertex to clypeus 2.09; frons width 1.87–2.10 (1.98); frons height 0.63–0.78 (0.71); prothoracic femur length 7.69–7.75 (7.72); mesothoracic femur length 9.92–10.23 (10.08); mesothoracic tibia length 7.69–7.84 (7.77); mesothoracic tarsus length 6.73–6.78 (6.76); metathoracic femur length 10.00; metathoracic tibia length 10.57–10.94 (10.75); metathoracic tarsus length 9.78–9.97 (9.88); pronotal elongation measure 0.29; pronotal shape measure 0.29–0.30 (0.29); head shape measure 0.40; frons shape measure 0.30–0.42 (0.36); anteroventral femoral spine count 14–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Pronotum (Fig. 48J): As described for males.

Prothoracic Legs: As described for males.

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, Liturgusa 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).

Figure 21.

Liturgusa dominica sp. n., ootheca: A dorsal B lateral.

Etymology.

A noun in apposition, Liturgusa dominica is named for the island of Dominica, where this species inhabits and appears to be endemic.

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA.

Type locality.

French Guiana: Kaw Mountain Res., Amazonas Lodge, 8-19 Feb 2005, Coll: K.B. Miller (Lat. 4.549389, Long. -52.213806).

Material examined.

Liturgusa milleri sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype French Guiana Kaw Mountain Res., Amazonas Lodge, 4°32'57.8"N, 52°12'49.7"W, 8-19 Feb 2005, Coll: K.B. Miller 4.549389, -52.213806 GSMC000264
Female Allotype French Guiana Kaw Mountain Res., Amazonas Lodge, 4°32'57.8"N, 52°12'49.7"W, 8-19 Feb 2005, Coll: K.B. Miller 4.549389, -52.213806 GSMC000260
3 Females Paratypes French Guiana Kaw Mountain Res., Amazonas Lodge, 4°32'57.8"N, 52°12'49.7"W, 8-19 Feb 2005, Coll: K.B. Miller 4.549389, -52.213806 GSMC000266, GSMC000314, GSMC003470
Male Paratype French Guiana Kaw Mountain Reserve, vicinity of Patawa, 30.IV/1.V.2011, 4.56888N, 52.21388W, elev. 282 m (at light), J. Rivera leg. 4.568880, -52.213880 GSMC003471
Female Paratype French Guiana St-Laurent du Maroni, Collection Le Moult, Coll. L. Chopard, 1919 5.487038, -54.008462 MNHN 017
Female Paratype French Guiana Arataye Affl Approuagues, Skm NE pied Saut Parare, 1-VI-1988, Chasse de nuit, L. Desutter & P. Grandcolas rec. 4.046724, -52.698087 MNHN 021
Female Paratype French Guiana Crique Venus, IX-1992, P. Peters 5.181326, -52.926674 MNHN 023
Female Paratype French Guiana Sinnamary, 3-6-VII-1977, Guyane, M. Descamps rec. 5.370512, -52.960320 MNHN 025
Male Paratype French Guiana St-Jean du Maroni, R. Benoist, 1914 5.487038, -54.008462 MNHN 035
Female Paratype French Guiana St-Laurent du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Juliet 5.487038, -54.008462 MNHN 052
Female Paratype French Guiana St-Jean du Maroni, R. Benoist, 1914, Mars 5.487038, -54.008462 MNHN 053
Female Paratype French Guiana Nouveau Chantier, collection Le Moult, Coll. L. Chopard, 1919, Mai MNHN 054
Male Paratype French Guiana Camopi, F. Geay - 1900 3.167700, -52.339455 MNHN 072
Male Paratype French Guiana Massikiri-Oyapock, 16-Nov-1969, Balachowski-Gruner, Oct.Nov. 1969 MNHN 073
Male Paratype French Guiana S. Boucher, 1984-85 MNHN 076
Female Paratype French Guiana NE, Route de Kaw, Caiman Camp env. 7.XII.2006, Snizek MSMC 003
Female Paratype French Guiana St-Laurent du Maroni 5.487038, -54.008462 OUMNH 009
Male Paratype French Guiana GSMC003015
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 Liturgusa milleri are restricted to the island of Dominica and Central America (Liturgusa dominica and Liturgusa cursor, respectively).

Description.

Male. (Fig. 22A) N=6: Body length 19.54–19.91 (19.67); forewing length 11.59–12.72 (12.07); hindwing length 9.26–10.19 (9.72); pronotum length 6.32–7.16 (6.69); prozone length 1.84–2.12 (1.95); pronotum width 1.90–2.10 (2.02); pronotum narrow width 1.27–1.39 (1.32); head width 4.42–4.74 (4.53); head vertex to clypeus 1.68–1.79 (1.72); frons width 1.46–1.63 (1.53); frons height 0.57–0.62 (0.59); prothoracic femur length 5.78–6.30 (5.92); mesothoracic femur length 7.96–8.70 (8.26); mesothoracic tibia length 6.11–7.09 (6.54); mesothoracic tarsus length 4.64–6.06 (5.58); metathoracic femur length 7.73–9.03 (8.46); metathoracic tibia length 8.22–9.55 (8.76); metathoracic tarsus length 8.80–9.35 (9.15); pronotal elongation measure 0.27–0.31 (0.29); pronotal shape measure 0.28–0.33 (0.30); head shape measure 0.38; frons shape measure 0.37–0.41 (0.39); anteroventral femoral spine count 13–15 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 22.

Liturgusa milleri sp. n., dorsal habitus: A holotype male from Kaw Mountain, French Guiana (CLEV GSMC000264) B allotype female from Kaw Mountain, French Guiana (CLEV GSMC000260).

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.

Female. (Fig. 22B) N=9: Body length 22.70–26.63 (25.22); forewing length 13.52–14.75 (14.15); hindwing length 9.65–11.33 (10.58); pronotum length 7.64–8.32 (7.97); prozone length 2.18–2.41 (2.32); pronotum width 2.36–2.60 (2.48); pronotum narrow width 1.43–1.75 (1.57); head width 5.29–5.66 (5.48); head vertex to clypeus 2.09–2.33 (2.18); frons width 1.94–2.12 (2.03); frons height 0.68–0.90 (0.77); prothoracic femur length 6.90–7.62 (7.28); mesothoracic femur length 9.07–9.82 (9.47); mesothoracic tibia length 7.24–8.13 (7.61); mesothoracic tarsus length 6.31–7.11 (6.75); metathoracic femur length 9.25–9.96 (9.67); metathoracic tibia length 9.89–10.75 (10.17); metathoracic tarsus length 9.46–10.65 (10.32); pronotal elongation measure 0.28–0.30 (0.29); pronotal shape measure 0.29–0.34 (0.31); head shape measure 0.38–0.41 (0.39); frons shape measure 0.34–0.42 (0.38); anteroventral femoral spine count 14–15 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Pronotum (Fig. 48L): As described for males.

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, Liturgusa 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, Table 1, Fig. 10; Henderson 1965: 206, 215.
Liturgusa actuosa: Terra 1995: 53; Jantsch 1999: 47; Ehrmann 2002: 206; Otte and Spearman 2005: 132; Agudelo et al. 2007: 116.
Type.

Holotype Male. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA, Type no. 5760.

Type locality.

Panama, Barro Colorado Island, Gatun Lake, Canal Zone, January 1, 1948. (Dr. Franz Schrader, no. 637.) (Lat. 9.164966, Long. -79.837098).

Material examined.

Liturgusa actuosa Rehn, 1950.

Sex Type Country Label Latitude Longitude Code
Male Holotype Panama Barro Colorado Island, Gatun Lake, Canal Zone, January 1, 1948. (Dr. Franz Schrader, no. 637.) 9.164966, -79.837098 ANSP
Male Paratype Panama Barro Colorado, C.Z., 4 I, 49, F. Schrader, Male, 657 9.164966, -79.837098 ANSP 040
Male Paratype Panama Barro Colorado Is. C.Z., F.-S.H. Schrader 9.164966, -79.837098 ANSP 041
Male Paratype Panama Barro Colorado, C.Z., 4 I, 49, F. Schrader, Male, 652 9.164966, -79.837098 ANSP 042
Female Paratype Panama Barro Colorado Is. C.Z., F.-S.H. Schrader 9.164966, -79.837098 ANSP 043
Male Paratype Panama Barro Colorado Is. C.Z., F.-S.H. Schrader 9.164966, -79.837098 ANSP 044
Male nontype Panama Barro Colorado I. C.Z. XI-22-44. Pres. by K.E. Frick Collector 9.164966, -79.837098 CAS 006
Female nontype Panama Barro Colorado I., C.Z., XI-22-44, Pres. by K.E. Frick Collector 9.164966, -79.837098 CAS 011
Male nontype Panama Barro Colorado I. C.Z. XI-22-44. Pres. by K.E. Frick Collector 9.164966, -79.837098 CAS 014
Male nontype Panama C.Z., Barro Colorado Is., 09°10'N, 79°50'W, 27-30 June 1973, Erwin & Hevel Central America Expedition, 1973 9.164966, -79.837098 USNM 012; USNM ENT 00873025
Male nontype Panama Barro Colorado I., Canal Zone, Panama, Jan. 1 1959, CNHM Panama, Zool. Exped. (1959) H.S. Dybas leg. 9.164966, -79.837098 FMNH 012
Nymph nontype Panama Barro Colorado, C.Z. Nov. 6 1930, H.T. Schwarz Coll. 9.164966, -79.837098 AMNH 008
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 Liturgusa annulipes rather than the sympatric Liturgusa cursor, but this study establishes that Liturgusa annulipes is not distributed in Central America and Liturgusa actuosa was being compared with an unknown taxon considered as Liturgusa 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 Liturgusa cameroni and Liturgusa 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 Liturgusa 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 Liturgusa cameroni and Liturgusa 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.

Redescription.

Male. (Fig. 23A) N=7: Body length 22.22–24.52 (23.50); forewing length 14.46–16.13 (15.44); hindwing length 11.63; pronotum length 6.59–7.08 (6.78); prozone length 1.92–2.12 (2.03); pronotum width 2.19–2.40 (2.28); pronotum narrow width 1.60–1.72 (1.64); head width 4.78–4.95 (4.88); head vertex to clypeus 1.87–2.00 (1.93); frons width 1.66–1.74 (1.70); frons height 0.61–0.73 (0.68); prothoracic femur length 6.10–6.57 (6.32); mesothoracic femur length 7.73–8.49 (8.02); mesothoracic tibia length 6.16–6.48 (6.32); mesothoracic tarsus length 5.44–5.54 (5.50); metathoracic femur length 7.78–8.93 (8.15); metathoracic tibia length 7.11–9.12 (8.59); metathoracic tarsus length 6.41–8.28 (7.71); pronotal elongation measure 0.29–0.31 (0.30); pronotal shape measure 0.33–0.35 (0.34); head shape measure 0.38–0.41 (0.40); frons shape measure 0.36–0.44 (0.40); anteroventral femoral spine count 13–15 (15); anteroventral tibial spine count 9–10 (10); posteroventral tibial spine count 7.

Figure 23.

Liturgusa actuosa Rehn, 1950, dorsal habitus: A male from Barro Colorado Island, Panama (USNM 012) B female from Barro Colorado Island, Panama (CAS 011).

Head (Fig. 44A): Juxta-ocular protuberances small, the apex in the middle 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 small cuticular mounds. Lateral ocelli oriented outward, a few degrees off perpendicular. Upper margin of clypeus convex, lower margin barely concave. Antennae pale basally fading gradually to dark brown or black near the middle. Broad black band extending straight over the medial carina of the frontal suture; lower region of frons with a broad black band; the clypeus mostly black with pale lateral and lower margins; the mandibles pale with lateral darkening; the labrum mostly dark; the vertex pale with black splotches and juxta-ocular protuberances mostly black anteriorly; the area around the ocelli mostly black. Palpi are pale.

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.

Redescription.

Female. (Fig. 23B) N=2: Body length 26.90–28.73 (27.81); forewing length 16.84–17.63 (17.23); hindwing length 14.55; pronotum length 7.79–7.97 (7.88); prozone length 2.37–2.46 (2.42); pronotum width 2.74–2.78 (2.76); pronotum narrow width 1.93–1.94 (1.94); head width 5.68–5.72 (5.70); head vertex to clypeus 2.46–2.51 (2.48); frons width 2.15; frons height 0.82–0.83 (0.82); prothoracic femur length 7.43–7.66 (7.54); mesothoracic femur length 8.43; mesothoracic tibia length 6.84; mesothoracic tarsus length 5.98; metathoracic femur length 8.43–8.62 (8.52); metathoracic tibia length 9.71–9.77 (9.74); metathoracic tarsus length 8.73; pronotal elongation measure 0.30–0.31 (0.31); pronotal shape measure 0.35; head shape measure 0.43–0.44 (0.43); frons shape measure 0.38; anteroventral femoral spine count 14–15 (14); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA.

Type locality.

Peru: Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson, Tissue 011, GSMC004007. (Lat. -3.62096, Long. -72.24744).

Material examined.

Liturgusa algorei sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 022 -3.620960, -72.247440 GSMC004007
Female Allotype Peru Loreto Province, Puerto Almendra -3.830525, -73.374000 GSMC004011
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 011 -3.620960, -72.247440 GSMC004000
Female Paratype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 029 -3.620960, -72.247440 GSMC004021
Female Paratype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 023 -3.620960, -72.247440 GSMC004023
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 001 -3.620960, -72.247440 GSMC004026
Male Paratype Peru Loreto Province, Puerto Almendra - Tissue 042 -3.830525, -73.374000 GSMC004029
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station -3.620960, -72.247440 GSMC004033
Male Paratype Equador Napo Rio Aguarico, San Pablo Tumba Emilio, VI-1985, K. Riede Rec. MNHN 037
Male Paratype Peru Loreto, Pebas, River Amazonas, -3.329066°S, -71.854168°E, 28 Feb 2010, Coll: J.J. Ramirez -3.329066, -71.854168 MNHN 095
Female Paratype Ecuador Napo: Yasuri Natl. Pk. 0°40'S, 76°20'W., Tiputini Biodiversity Station, 14-19 FEB 1998. UofT Field Course, per DC Darling. ROM 980000 -0.637742, -76.150216 ROM 001
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 Liturgusa 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 Liturgusa 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 Liturgusa krattorum.

Description.

Male. (Fig. 24A) N=7: Body length 22.00–24.36 (23.42); forewing length 13.17–14.98 (14.31); hindwing length 10.89–11.64 (11.28); pronotum length 7.11–7.79 (7.52); prozone length 1.94–2.18 (2.08); pronotum width 2.15–2.33 (2.23); pronotum narrow width 1.52–1.68 (1.57); head width 4.51–5.10 (4.83); head vertex to clypeus 1.89–2.03 (1.97); frons width 1.69–1.77 (1.72); frons height 0.63–0.75 (0.69); prothoracic femur length 6.20–7.00 (6.70); mesothoracic femur length 7.07–10.52 (9.43); mesothoracic tibia length 5.67–8.22 (7.37); mesothoracic tarsus length 5.36–6.94 (6.36); metathoracic femur length 8.97–10.71 (9.91); metathoracic tibia length 9.06–10.85 (9.88); metathoracic tarsus length 9.77–11.08 (10.44); pronotal elongation measure 0.27–0.28 (0.28); pronotal shape measure 0.28–0.31 (0.30); head shape measure 0.39–0.44 (0.41); frons shape measure 0.36–0.44 (0.40); anteroventral femoral spine count 15–16 (15); anteroventral tibial spine count 10–11 (10); posteroventral tibial spine count 7.

Figure 24.

Liturgusa algorei sp. n., dorsal habitus: A holotype male from Loreto, Peru (CLEV GSMC004007) B allotype female from Loreto, Peru (CLEV GSMC004011).

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.

Meso- and Metathoracic Legs: Femora with faint ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment equal to remaining segments combined.

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.

Female. (Fig. 24B) N=4: Body length 31.26–32.15 (31.68); forewing length 17.99–19.15 (18.61); hindwing length 14.78–15.39 (15.08); pronotum length 9.11–9.99 (9.74); prozone length 2.70–2.88 (2.81); pronotum width 2.94–3.14 (3.07); pronotum narrow width 1.84–2.08 (1.97); head width 5.75–6.28 (6.12); head vertex to clypeus 2.60–2.76 (2.69); frons width 2.31–2.54 (2.43); frons height 0.89–0.96 (0.92); prothoracic femur length 8.75–9.23 (8.99); mesothoracic femur length 11.24–12.09 (11.81); mesothoracic tibia length 8.58–9.39 (9.13); mesothoracic tarsus length 7.73–8.22 (7.95); metathoracic femur length 11.34–12.45 (12.01); metathoracic tibia length 11.62–12.49 (12.20); metathoracic tarsus length 10.81–12.5 (11.95); pronotal elongation measure 0.28–0.30 (0.29); pronotal shape measure 0.29–0.34 (0.32); head shape measure 0.42–0.47 (0.44); frons shape measure 0.37–0.39 (0.38); anteroventral femoral spine count 14–15 (15); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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.

Meso- and Metathoracic Legs: Femora with ventral (posterior) and dorsal (anterior) carina present.

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, Liturgusa 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.

Type.

Holotype Male, pinned. American Museum of Natural History, New York, NY, USA.

Type locality.

Venezuela: Rancho Grande, nr. Maracay, Ven. 17-VI-1948 (Lat. 10.350000, Long. -67.683330).

Material examined.

Liturgusa cameroni sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Guyana Kartabo, Bartica District, 1921 6.242050, -59.306552 ANSP 047
Female Allotype Venezuela Estado de Aragua, Rancho Grande, N. of Maracay, 1300 m, 20.II.1987, Edward S. Ross 10.350000, -67.683330 CAS 003
Male Paratype Venezuela Rancho Grande, nr. Maracay, 17-VI-1948 10.350000, -67.683330 AMNH 016
Male Paratype Venezuela Rancho Grande, nr. Maracay, 23-VI-1946 10.350000, -67.683330 AMNH 017
Female Paratype Venezuela San Esteban, X-XI 1910, M.A. Carriker Jr. 10.422557, -68.012585 ANSP 019
Male Paratype Guyana Kartabo, Bartica District, 1921 6.242050, -59.306552 ANSP 045
Female Paratype Venezuela San Rafael, Cumanacoa, Sucre, G. Netting, Nov 29 1929 10.227884, -63.947462 ANSP 114
Female Paratype Venezuela La Guaira, Robinson, S.H. Scudder Collection 10.596235, -66.934786 ANSP 117
Female Paratype Venezuela La Guaira, Robinson, S.H. Scudder Collection 10.596235, -66.934786 ANSP 118
Male Paratype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 075
Female Paratype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 078
Female Paratype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 079
Female Paratype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 080
Male Paratype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 085
Nymph nontype Venezuela Petare, 880 mtrs., 21.V.1926, H.E. Box Collector 10.500706, -66.799504 BMNH 077
Diagnosis.

Nearly identical to Liturgusa 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 Liturgusa 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 Liturgusa nubeculosa.

Description.

Male. (Fig. 25A) N=5: Body length 21.88–26.52 (24.66); forewing length 13.40–16.34 (15.19); pronotum length 7.21–8.74 (8.09); prozone length 2.08–2.49 (2.31); pronotum width 2.23–2.68 (2.48); pronotum narrow width 1.73–2.01 (1.88); head width 4.87–5.54 (5.23); head vertex to clypeus 1.80–2.12 (1.97); frons width 1.76–2.00 (1.89); frons height 0.55–0.73 (0.65); prothoracic femur length 6.44–8.10 (7.29); mesothoracic femur length 8.54–10.93 (9.89); mesothoracic tibia length 6.66–8.49 (7.96); mesothoracic tarsus length 5.57–7.51 (6.76); metathoracic femur length 7.27–11.64 (9.78); metathoracic tibia length 8.97–11.76 (10.55); metathoracic tarsus length 10.66; pronotal elongation measure 0.28–0.29 (0.29); pronotal shape measure 0.30–0.32 (0.31); head shape measure 0.36–0.39 (0.38); frons shape measure 0.30–0.38 (0.34); anteroventral femoral spine count 15–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 25.

Liturgusa cameroni sp. n., dorsal habitus: A holotype male from Guyana (ANSP 047) B allotype female from Maracay, Venezuela (CAS 003).

Head (Fig. 44E): Transverse, juxta-ocular protuberances small, the apex in the lateral third; the vertex between the parietal sutures is straight and broad; vertex even or slightly below the dorsal margin of the eyes. Frontal suture with a medial carina forming an arc the middle with an obtuse angle. Ocelli small and protruding on small cuticular mounds. Lateral ocelli oriented outward. Frons highly transverse, the region below the antennal insertions very narrow; lower medial region with black marking. Clypeus transverse; upper margin straight, lower margin concave. Antennae pale basally, gradually fading to black. Upper region of clypeus pale, lower half brown; mandibles pale or brown; the vertex and juxta-ocular protuberances splotched with brown; the area adjacent to lateral ocelli black. Palpi are pale.

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.

Female. (Fig. 25B) N=6: Body length 27.44–35.27 (31.21); forewing length 15.35–19.92 (17.30); hindwing length 14.68; pronotum length 9.07–11.12 (10.01); prozone length 2.59–3.17 (2.83); pronotum width 3.00–3.47 (3.23); pronotum narrow width 2.19–2.43 (2.27); head width 6.10–6.86 (6.48); head vertex to clypeus 2.48–2.78 (2.63); frons width 2.36–2.61 (2.46); frons height 0.78–0.97 (0.86); prothoracic femur length 8.02–10.43 (9.04); mesothoracic femur length 9.81–12.73 (11.14); mesothoracic tibia length 7.96–10.25 (8.94); mesothoracic tarsus length 6.96–8.69 (7.63); metathoracic femur length 9.70–12.13 (10.83); metathoracic tibia length 10.54–14.05 (12.09); metathoracic tarsus length 9.90–12.31 (10.99); pronotal elongation measure 0.28–0.29 (0.28); pronotal shape measure 0.31–0.34 (0.32); head shape measure 0.40–0.42 (0.41); frons shape measure 0.33–0.39 (0.35); anteroventral femoral spine count 16–17 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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, Liturgusa cameroni is named for Stephen L. Cameron for his contributions to Mantodea sampling and his valuable collaboration.

Type.

Holotype Male, pinned. Cleveland Museum of Natural History, Cleveland, OH, USA.

Type locality.

Peru: Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson, Tissue 005, GSMC004004. (Lat. -3.62096, Long. -72.24744).

Material examined.

Liturgusa krattorum sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10–17 February 2013, Coll: G.J. Svenson - Tissue 005 -3.620960, -72.247440 GSMC004004
Female Allotype Peru Loreto Province, Puerto Almendra - Tissue 040 -3.830525, -73.374000 GSMC004032
Female Paratype Peru Rio Maranon, XII-30, F 6079, H. Bassler Collection -4.832821, -76.639033 AMNH 003
Male Paratype Peru Monson Valley, Tingo Maria, X-19-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 008
Female Paratype Peru 20 mi. W. of Pucalipa, X-3-1954, E.I. Schlinger & E.S. Ross collectors -8.354749, -74.552113 CAS 019
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson - Tissue 026 -3.620960, -72.247440 GSMC004001
Male Paratype Peru Loreto Province, Puerto Almendra - Tissue 033 -3.830525, -73.374000 GSMC004018
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson - Tissue 002 -3.620960, -72.247440 GSMC004034
Male Paratype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson - Tissue 009 -3.620960, -72.247440 GSMC004035
Male Paratype Peru Dept. Loreto Rio Yubineto, 1-VII - 1.VIII - 1978, M. Descamps rec. -0.999780, -74.253238 MNHN 024
Female Paratype Colombia Dept. Amazonas, 30 KM. Aval de la Chorrera, Rio Igara - Parana, 1.2.II.1974, J. Desplats Rec. -0.754819, -73.007213 MNHN 082
Female Paratype Peru Dept. Loreto Rio Yubineto, 1-VII - 1.VIII - 1978, M. Descamps rec. -0.999780, -74.253238 MNHN 084
Male Paratype Peru Loreto, Pebas, River Amazonas, -3.329066°S, -71.854168°E, 28 Feb 2010, Coll: J.J. Ramirez -3.329066, -71.854168 MNHN 093
Female Paratype Peru Loreto, Iquitos, Nov. 2010, Collection Stiewe -3.741872, -73.272190 MSMC 002
Male Paratype Ecuador Pastaza; Ashuara, Rio Macuma, 10km from Rio Morona, 300m, VII:7-16:1971, le. B. Malkin -2.753512, -77.444899 FMNH 004
Female Paratype Ecuador Pastaza; Ashuara, Rio Macuma, 10km from Rio Morona, 300m, VII:7-16:1971, le. B. Malkin -2.753512, -77.444899 FMNH 011
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 Liturgusa 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 Liturgusa purus and Liturgusa 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 Liturgusa purus, but more rusty than Liturgusa 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 Liturgusa algorei.

Description.

Male. (Fig. 26A) N=8: Body length 23.24–24.68 (23.94); forewing length 14.39–15.50 (14.91); hindwing length 11.33–12.48 (11.81); pronotum length 7.27–7.85 (7.62); prozone length 2.06–2.24 (2.13); pronotum width 2.21–2.41 (2.33); pronotum narrow width 1.51–1.71 (1.59); head width 4.70–4.92 (4.85); head vertex to clypeus 1.88–2.03 (1.96); frons width 1.64–1.77 (1.70); frons height 0.63–0.73 (0.67); prothoracic femur length 6.62–7.09 (6.90); mesothoracic femur length 9.40–10.17 (9.79); mesothoracic tibia length 7.46–8.28 (7.95); mesothoracic tarsus length 6.69–7.45 (7.06); metathoracic femur length 9.95–10.80 (10.28); metathoracic tibia length 9.93–10.86 (10.50); metathoracic tarsus length 10.12–11.52 (11.00); pronotal elongation measure 0.27–0.29 (0.28); pronotal shape measure 0.29–0.32 (0.30); head shape measure 0.40–0.42 (0.40); frons shape measure 0.36–0.42 (0.40); anteroventral femoral spine count 15–17 (16); anteroventral tibial spine count 10–11 (10); posteroventral tibial spine count 7.

Figure 26.

Liturgusa krattorum sp. n., dorsal habitus: A holotype male from Loreto, Peru (CLEV GSMC004004) B allotype female from Loreto, Peru (CLEV GSMC004032).

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. Posterior 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.

Meso- and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment approximately equal to remaining segments combined.

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.

Female. (Fig. 26B) N=6: Body length 30.58–37.69 (33.14); forewing length 17.74–19.93 (18.91); hindwing length 15.02–15.97 (15.35); pronotum length 9.32–10.93 (9.86); prozone length 2.66–3.00 (2.78); pronotum width 2.94–3.40 (3.10); pronotum narrow width 2.08–2.24 (2.15); head width 5.90– 6.84 (6.22); head vertex to clypeus 2.57–2.91 (2.72); frons width 2.32–2.61 (2.42); frons height 0.86–0.97 (0.92); prothoracic femur length 8.37–9.90 (8.94); mesothoracic femur length 11.25–12.67 (11.80); mesothoracic tibia length 9.08–10.45 (9.65); mesothoracic tarsus length 7.59–8.47 (7.95); metathoracic femur length 11.44–13.44 (12.20); metathoracic tibia length 11.53–14.03 (12.91); metathoracic tarsus length 11.71–14.80 (13.13); pronotal elongation measure 0.28–0.29 (0.28); pronotal shape measure 0.30–0.33 (0.32); head shape measure 0.42–0.45 (0.44); frons shape measure 0.35–0.40 (0.38); anteroventral femoral spine count 14–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Head (Fig. 44H): Juxta-ocular protuberances large, the apex in the medial third; vertex well above the dorsal margin of the eyes. Upper margin of clypeus barely convex, lower margin straight with medial bulge.

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.

Meso- and Metathoracic Legs: Femora with ventral (posterior) carina; dorsal (anterior) carina obvious.

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, Liturgusa 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.

Type.

Holotype Female, pinned. Muséum national d’Histoire naturelle, Paris, France.

Type locality.

French Guiana: Guyane Franc, St-Laurent du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Octobre (Lat. 5.487038, Long. -54.008462).

Material examined.

Liturgusa maroni sp. n.

Sex Type Country Label Latitude Longitude Code
Female Holotype French Guiana St-Laurent du Maroni, Collection Le Moult, Coll. L. Chopard, 1919, Octobre 5.487038, -54.008462 MNHN 019
Female Paratype French Guiana Petit Saut, 8-II-1994, P.E. Rouland 5.069416, -53.047566 MNHN 061
Diagnosis.

A medium size species known only from female specimens from French Guiana. Most similar to Liturgusa 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.

Description.

Female. (Fig. 7B) N=2: Body length 27.81–28.75 (28.28); forewing length 15.94–16.24 (16.09); hindwing length 12.69–12.78 (12.73); pronotum length 8.98–9.19 (9.09); prozone length 2.54–2.63 (2.58); pronotum width 3.00–3.04 (3.02); pronotum narrow width 2.01–2.21 (2.11); head width 6.18–6.29 (6.23); head vertex to clypeus 2.49–2.66 (2.58); frons width 2.27–2.30 (2.28); frons height 0.78; prothoracic femur length 8.49–8.88 (8.68); mesothoracic femur length 11.09–11.74 (11.41); mesothoracic tibia length 8.95–9.36 (9.15); mesothoracic tarsus length 7.78–8.05 (7.92); metathoracic femur length 11.25–11.77 (11.51); metathoracic tibia length 12.39–12.82 (12.60); metathoracic tarsus length 12.11–12.37 (12.24); pronotal elongation measure 0.28–0.29 (0.28); pronotal shape measure 0.33; head shape measure 0.40–0.43 (0.41); frons shape measure 0.34; anteroventral femoral spine count 16; anteroventral tibial spine count 10; posteroventral tibial spine count 7.

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, Liturgusa maroni is named for the Maroni River near the type locality of Saint Laurent du Maroni on the border of French Guiana and Suriname.

Liturgusa nubeculosa Gerstaecker, 1889

http://species-id.net/wiki/Liturgusa_nubeculosa

Liturgusa nubeculosa: Gerstaecker 1889: 54–56; Bertkau 1889: 87; Giglio-Tos 1927: 294; Beier 1935: 11; Terra 1995: 54; Jantsch 1999: 48; Lombardo and Agabiti 2001: 90, 97; Ehrmann 2002: 207; Otte and Spearman 2005: 133; Agudelo et al. 2007: 116, 142.
Liturgousa nubeculosa: Westwood 1889: 5, 51; Rehn 1935: 199; Rehn 1954: 177, pl. 1, fig. 2; Cerdá 1996: 76, Fig. 1.
Hagiomantis nubeculosa: Kirby 1904: 271.
= Liturgusa peruviana: Giglio-Tos 1914: 77; Giglio-Tos 1927: 293; Beier 1935: 11; Rehn 1954: 177; Marshall 1975: 322; Terra 1995: 54; Ehrmann 2002: 207; Otte and Spearman 2005: 133. syn. n. (Rehn 1935)
Type.

Holotype Female. Ernst-Moritz-Arndt-Universität Greifswald, Germany.

Type locality.

Brazil: Ega, Fonteboa (Amazonas) (Lat. -2.522585, Long. -66.097224).

Material examined.

Liturgusa nubeculosa Gerstaecker, 1889.

Sex Type Country Label Latitude Longitude Code
Female Holotype Brazil Ega, Fonteboa (Amazonas) -2.522585, -66.097224 EMAU
Female Holotype (peruviana) Peru Palcaru, 1904-65 BMNH
Female nontype Peru Middle Rio Ucayali, II -27, F 6085, H. Bassler Collection Acc. 33591 -9.823201, -73.960930 AMNH 001
Female nontype Venezuela Amazonas: Rio Mavaca Cp. 65°06'W, 2°2'N, 150m. III-16/27-1989, David Grimaldi, Exp. Phipps-Fudeci 2.033333, -65.100000 AMNH 025
Female nontype Ecuador Pastaza: Cuisimi, on Rio Cuisimi 150km. SE Puyo, 350 m. June 1–5, 1971, B. Malkin -2.404277, -77.040975 AMNH 026
Female nontype Brazil Obidos, Para, Brazil, VIII 23 1919 (H.S. Parish) -1.899285, -55.528371 ANSP 018
Female nontype Peru Leonpampa 110 k. E. Huanaco, Huanaco Prov. Peru, Tropical Jungle, December 1937, Felix Woytkowski ANSP 039
Male nontype Brazil Manacapuru, S.M. Klages, March 1926 -3.286094, -60.640173 ANSP 106
Female nontype Brazil Nova Oiinda, Rio Purus, S.M. Klages, May 1922, Carn. Mus. 6962 ANSP 107
Male nontype Brazil Para BMNH 004
Female nontype Peru Yuras, 67 mi. E. of Tingo Maria. 350 m. XII-11-54. E.I. Schlinger & E.S. Ross collectors -9.231743, -76.321433 CAS 016
Female nontype Peru Monzon Valley, Tingo Maria, IX-23-1954, E.I. Schlinger & E.S. Ross collectors -9.314153, -76.006745 CAS 018
Female nontype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000269
Female nontype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3–9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000270
Female nontype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3–9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000273
Female nontype Bolivia Dpto. Santa Cruz, Reserva Natural Potrerillo del Guenda, 17°40.281'S, 063°27.451'W, 400 m, 3-9.XI.2009, at MV.UV lights & gen. coll., Coll: G.J. Svenson -17.671350, -63.457517 GSMC000308
Female nontype Peru Loreto, Pacaya Samiria, Cocha Shinguito; TLE & MGP; TSnl Insecticidal fog of Fog Tree 2 GSMC003072
Male nontype Peru Loreto Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson -3.620960, -72.247440 GSMC004006
Male nontype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 024 -3.620960, -72.247440 GSMC004006
Female nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004010
Female nontype Peru Loreto Province, Puerto Almendra - Tissue 035 -3.830525, -73.374000 GSMC004010
Male nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004012
Male nontype Peru Loreto Province, Puerto Almendra - Tissue 044 -3.830525, -73.374000 GSMC004012
Male nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004013
Male nontype Peru Loreto Province, Puerto Almendra -3.830525, -73.374000 GSMC004013
Male nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004014
Male nontype Peru Loreto Province, Puerto Almendra - Tissue 043 -3.830525, -73.374000 GSMC004014
Female nontype Peru Loretro Province, Madre Selva Biological Research Station, -3.62096, -72.24744, 10-17 February 2013, Coll: G.J. Svenson -3.620960, -72.247440 GSMC004022
Female nontype Peru Loreto Province, Madre Selva Biological Research Station - Tissue 021 -3.620960, -72.247440 GSMC004022
Nymph nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004030
Female nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004031
Female nontype Peru Loreto Province, Puerto Almendra - Tissue 039 -3.830525, -73.374000 GSMC004031
Nymph nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004040
Nymph nontype Peru Loreto Province, Puerto Almendra, -3.830525, -73.374, 108 m, 19-21 February 2013, Coll: G.J. Svenson, Tissue 032 -3.830525, -73.374000 GSMC004045
Male nontype Peru Madre de Dios, Rio Tambopata Res. 30km (air) sw Pto. Maldonato, 290m, 12°50'S, 069°20'W, Smithsonian Institution Canopy Fogging Project, T.L. Erwin et al. colls. 30 Apr84, 03/O&S (01), Fogging 00017348 -12.833333, -69.333333 GSMC004055
Male nontype Colombia Dept. Amazonas, Rio Igara Parana, 30 km aval La Chorrera, VI-VIII 1974, M. Descamps rec. -1.197385, -72.937475 MNHN 009
Female nontype Ecuador Limoncocha, 25-VII-1983, K. Riede rec. -0.400848, -76.618481 MNHN 056
Female nontype Peru Dept. Loreto Colonia, Amont Conflt. Rios Zumun & Yahuasyacu, 20-V-20-VI-1978, M. Descamps rec. MNHN 057
Female nontype Brazil Benjamin Constant, IX 1979, AM, B. Silva rec., C. Seabia leg. -4.383010, -70.042251 MNHN 058
Female nontype Peru Loreto, Maynas, Picuroyacu, apres Sta Clotilde, N. Iquitos, 130 mts., 10.II.2010, S03.37.04 - W 73.15.44, M. Dottax leg. -3.617778, -73.262222 MNHN 089
Female nontype Peru Loreto, Picuroyacu, Jan 2010, Coll: J.J. Ramirez -3.617778, -73.262222 MNHN 094
Male nontype Peru Loreto, Maynas, Picuroyacu, apres Sta Clotilde, N. Iquitos, 130 mts., 10.II.2010, S03.37.04 - W 73.15.44, M. Dottax leg. -3.617778, -73.262222 MNHN 098
Male nontype Peru Loreto, Maynas, Picuroyacu, apres Sta Clotilde, N. Iquitos, 130 mts., 10.II.2010, S03.37.04 - W 73.15.44, M. Dottax leg. -3.617778, -73.262222 MNHN 099
Female nontype Ecuador Pr. Napo, Rio Tiputini, Pindo, Mandaripanga, 76°44'W, 0°43'S, 30 IX / 15 × 1997, Amedegnato / Poulain rec. -0.716667, -76.733333 MNHN 204
Female nontype Ecuador Pr. Napo, Rio Tiputini, Pindo, Mandaripanga, 76°44'W, 0°43'S, 30 IX / 15 × 1997, Amedegnato / Poulain rec. -0.716667, -76.733333 MNHN 205
Female nontype Bolivia Dpto. Santa Cruz, Prov. Andres Ibanez, Loc. Espejillos, 30-VIII-1990, col. Ma. Estker Moutano -17.718841, -63.438001 MNKM 001
Female nontype Peru Loreto, Iquitos, Nov. 2010, Collection Stiewe -3.741872, -73.272190 MSMC 001
Female nontype Peru Chamicuros, Bartlett OUMNH 007
Female nontype Brazil Para OUMNH 017
Male nontype Brazil Amazonas, Rio Janauaca, 40 km SW Manaus, 10 Mar 1979, 03°20'S, 060°17'W, White water innundation forest canopy fogged with Pyrethrin Sample #57, Montgomery, Erwin, Schimmel, Krischik, Date, Bacon colls. -3.333333, -60.283333 USNM 005; USNM ENT 00873027
Male nontype Brazil Amazonas, Rio Janauaca, 40 km SW Manaus, 10 Mar 1979, 03°20'S, 060°17'W, White water innundation forest canopy fogged with Pyrethrin Sample #52, Montgomery, Erwin, Schimmel, Krischik, Date, Bacon colls. -3.333333, -60.283333 USNM 009; USNM ENT 00873026
Female nontype Ecuador N.P., Sant Cecilia, III-25-31-1969, P. & P. Spangler USNM 031; USNM ENT 00873028
Female nontype Colombia Amazonas, Rio Cotuhe, St. Lucia + Tarapaca, 4 Feb. Megers -2.889475, -69.745012 USNM 032; USNM ENT 00873029
Male nontype Ecuador Road between El Puyo & Puerto Napo, VII-1964, Dr. Ch. Gregoire -1.229112, -77.879766 USNM 068; USNM ENT 00873030
Female nontype Peru Nord-Peru, Mishuyacu (Maranon-Gebiet) -5.910456, -76.103566 ZMHB 006
Female nontype Ecuador Canálos, S.V. Feyer -1.589524, -77.746701 ZMHB 008
Female nontype Brazil Obidos, Amazonas, I. Michaelis leg., vend. 13.IX 1900 -1.899285, -55.528371 ZMUH 001
Male nontype Ecuador Pastaza; 300m. Rio Macuma, 10km from Rio Marona, VII:5-7:1971, leg. B. Malkin -2.753512, -77.444899 FMNH 003
Male nontype Ecuador Pastaza; Cusuimi, Rio Cusuimi, 150km SE of Puyo, V:15-31:1971, leg. B. Malkin -2.391736, -77.047683 FMNH 006
Taxonomic history.

The species was described early relative to others in the genus, but was only included in taxon lists without receiving revisionary attention. In 1904 Kirby moved the species to Hagiomantis, but this action was ignored by other taxonomists since Giglio-Tos included the species within Liturgusa along with all subsequent works.

In 1954, James Rehn synonymized Liturgusa peruviana Giglio-Tos, 1914, with its senior synonym, Liturgusa 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 Liturgusa 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 Liturgusa 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 Liturgusa cameroni.

Description.

Male. (Fig. 27A) N=11: Body length 24.64–27.05 (25.62); forewing length 15.12–16.26 (15.73); hindwing length 11.94–12.67 (12.37); pronotum length 7.39–8.84 (8.06); prozone length 2.03–2.34 (2.19); pronotum width 2.42–2.68 (2.56); pronotum narrow width 1.74–2.06 (1.86); head width 4.79–5.47 (5.27); head vertex to clypeus 1.90–2.15 (2.07); frons width 1.77–2.02 (1.91); frons height 0.61–0.79 (0.70); prothoracic femur length 6.63–7.51 (7.14); mesothoracic femur length 8.93–10.67 (9.70); mesothoracic tibia length 7.03–8.34 (7.59); mesothoracic tarsus length 6.30–7.43 (6.64); metathoracic femur length 9.01–10.30 (9.69); metathoracic tibia length 9.60–10.89 (10.34); metathoracic tarsus length 9.06–10.84 (9.80); pronotal elongation measure 0.26–0.28 (0.27); pronotal shape measure 0.30–0.33 (0.32); head shape measure 0.38–0.41 (0.39); frons shape measure 0.34–0.40 (0.37); anteroventral femoral spine count 15–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 27.

Liturgusa nubeculosa Gerstaecker, 1889, dorsal habitus: A male from Loreto, Peru (CLEV GSMC004012) B female from Santa Cruz, Bolivia (CLEV GSMC000263).

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, 27B, 28A, 28C) N=23: Body length 32.17–52.03 (37.48); forewing length 19.32–26.96 (21.76); hindwing length 15.98–19.15 (17.44); pronotum length 9.77–13.12 (10.85); prozone length 2.76–3.69 (3.04); pronotum width 3.16–4.49 (3.62); pronotum narrow width 2.09–3.07 (2.54); head width 6.37–7.91 (6.86); head vertex to clypeus 2.58–3.38 (2.90); frons width 2.46–3.23 (2.70); frons height 0.89–1.23 (1.00); prothoracic femur length 8.54–12.24 (9.88); mesothoracic femur length 10.74–14.62 (12.30); mesothoracic tibia length 8.84–12.35 (9.94); mesothoracic tarsus length 7.71–10.74 (8.66); metathoracic femur length 10.88–14.87 (12.26); metathoracic tibia length 12.22–17.08 (13.55); metathoracic tarsus length 10.68–13.38 (12.34); pronotal elongation measure 0.27–0.29 (0.28); pronotal shape measure 0.32–0.35 (0.33); head shape measure 0.40–0.44 (0.42); frons shape measure 0.34–0.42 (0.37); anteroventral femoral spine count 15–17 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7 (one female with 8 on left tibia).

Figure 28.

Liturgusa nubeculosa Gerstaecker, 1889, and Liturgusa peruviana Giglio-Tos, 1914 syn. n., dorsal habitus and labels. Liturgusa nubeculosa: A holotype female (EMAU) B labels. Liturgusa peruviana: C holotype female (BMNH) D labels.

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.

Pronotum (Fig. 49L): Prozone elongate with near parallel lateral margins before tapering anteriorly; the margins with few small tubercles.

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.

Type.

Holotype Male, pinned. Academy of Natural Sciences of Drexel University, Philadelphia, PA, USA.

Type locality.

Brazil, Hyutanahan, Rio Purus, March 1922, S.M. Klages (Lat. -5.602502, Long. -63.221263).

Material examined.

Liturgusa purus sp. n.

Sex Type Country Label Latitude Longitude Code
Male Holotype Brazil Hyutanahan, Rio Purus, Klages, S.M., March 1922 -5.602502, -63.221263 ANSP 101
Female Allotype Brazil Hyutanahan, Rio Purus, Klages, S.M., March 1922 -5.602502, -63.221263 ANSP 103
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 Liturgusa krattorum and Liturgusa 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, Liturgusa purus has a posterior prothoracic femoral genicular spine that originates well proximal to the beginning of the genicular lobe, which is unique among Liturgusa krattorum, Liturgusa algorei, and other Cursor Group species.

Description.

Male. (Fig. 29A) N=2: Body length 29.16; forewing length 15.12–17.65 (16.39); hindwing length 11.86; pronotum length 7.42; prozone length 2.00; pronotum width 2.36; pronotum narrow width 1.61; head width 4.90; head vertex to clypeus 2.03; frons width 1.71; frons height 0.68; prothoracic femur length 6.64; mesothoracic femur length 9.40–11.15 (10.28); mesothoracic tibia length 7.44–8.83 (8.14); mesothoracic tarsus length 6.89; metathoracic femur length 9.67–11.48 (10.58); metathoracic tibia length 9.99–12.00 (10.99); metathoracic tarsus length 10.48–11.13 (10.80); pronotal elongation measure 0.27; pronotal shape measure 0.32; head shape measure 0.41; frons shape measure 0.40; anteroventral femoral spine count 15–16 (16); anteroventral tibial spine count 10; posteroventral tibial spine count 7.

Figure 29.

Liturgusa purus sp. n., dorsal habitus: A holotype male from Rio Purus, Brazil (ANSP 101) B allotype female from Rio Purus (ANSP 103).

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.

Meso- and Metathoracic Legs: Femora with faint ventral (posterior) carina; dorsal (anterior) carina faint. Mesotarsi with first segment equal to remaining segments combined.

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.

Pronotum: Elongate with a moderately defined supra-coxal bulge. Metazone with concave lateral margins; margins with numerous small tubercles; posterior margin medially emarginate.

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.

Wings: Forewings mottled with brown, green, and pale coloration.

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 projections in the distal half of the abdomen. Supra-anal plate slightly transverse, tapering to a rounded, emarginate terminus.

Etymology.

A noun in apposition, Liturgusa purus is named for Rio Purus, Brazil.

Fuga gen. n.

http://zoobank.org/43C4414B-EAF2-473B-BD34-8DAB040D3699

http://species-id.net/wiki/Fuga

Mantis (partim): Audinet Serville 1838: 199.
Oxypilus (partim): De Haan 1842: 84.
Liturgousa (partim): Saussure 1869: 62; Brauer 1870: 92; Saussure 1871b: 102 (♀ only; partim); Westwood 1889: 4, 49, pl. 2, fig. 3; Kirby 1904: 271; Werner 1906: 372; Chopard 1911: 323; Chopard 1916: 164; Rehn 1935: 199, pl. 8, fig. 4; Hughes-Schrader 1943: 266, 280, Table 1, Figs 19–28; Hughes-Schrader 1948: 267; Hughes-Schrader 1950: 11, Table 1; Cerdá 1996: 75.
Liturgusa (partim): Hebard 1922: 337; Giglio-Tos 1927: 294; Beier 1935: 11; Terra 1995: 53, Figs 85–87; Jantsch 1999: 48; Ehrmann 2002: 206; Otte and Spearman 2005: 132; Agudelo et al. 2007: 116, 141.
Hagiomantis (partim): Piza 1965: 130; Piza 1966: 8; Terra 1995: 54; Ehrmann 2002: 163–164; Otte and Spearman 2005: 129–130.
Type species.

Mantis