Liturgusa Saussure, 1869

Svenson, Gavin J., 2014, Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini), ZooKeys 390, pp. 1-214 : 14-23

Liturgusa Saussure, 1869

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, 21 A–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

Clave Para las Especies (translation by Julio Rivera)