Three new species of the genus Ripipteryx from Colombia (Orthoptera, Ripipterygidae)

Abstract Three new species of Ripipteryx Newman (Orthoptera: Tridactyloidea: Ripipterygidae) are described from Colombia; namely Ripipteryx diegoi sp. n. (Forceps Group) and Ripipteryx guacharoensis sp. n. (Marginipennis Group) from Parque Nacional Natural Cueva de los Guacharos in Huila, and Ripipteryx gorgonaensis sp. n. (Crassicornis Group) from Parque Nacional Natural Gorgona in Cauca. Ripipteryx diegoi sp. n. is characterized by the antennae black with white spots on flagellomeres 3–7, male subgenital plate with median ridge forming a bilobed setose process, epiproct produced laterally near its base and phallic complex with virga thickened distally and not reaching beyond the membrane. Ripipteryx guacharoensis sp. n. is characterized by the antennae thick with white spots present dorsally on flagellomeres 1–4 and 8, epiproct narrow and triangular, uncus reduced and lacking a distal hook, phallic complex with a concave ventral plate and a dorsal elevation in the middle extended to the virga, and the virga itself with two small projections basally. Ripipteryx gorgonaensis sp. n. is characterized by the epiproct with a lateral notch, antennae with a white dorsal spot on flagellomere 1 and flagellomeres 4–7 entirely white. The antennal color pattern of Ripipteryx gorgonaensis sp. n. strongly resembles that of Ripipteryx atra but differs from the latter in the absence of any significant morphological modification of the flagellomeres.


Introduction
Ripipteryx Newman, 1834, or mud crickets (Orthoptera: Tridactyloidea: Ripipterygidae), comprises some 45 species of small, dark-colored, cricket-like orthopterans usually found near rivers, in bare soil, and in the moist zones of gallery forests. Like many of their relatives in the larger cosmopolitan family Tridactylidae, the mud crickets are able to jump from the surface of water. The genus is readily distinguished from Mirhipipteryx, the only other genus in the family, by its comparatively larger size (body 5.5-14.0 mm long), interocular distance at least half the width of the compound eyes, metatarsus approximately equal in length to the metatibial spurs, and the distinctly sclerotized lateral valvulae of the phallus (Günther 1969;Heads 2010). Species of the genus are usually black or very dark brown, often with contrasting white, yellow and occasionally red markings (Heads 2010). Some species are a dark metallic blue in life, though this coloration often fades to brown or black after death. While ripipterygids are common in many habitats throughout the Neotropics, they are often overlooked by collectors due both to their small size and their fast and very active movements making it difficult to secure specimens. In addition to the paucity of specimens in collections, chronic under-sampling and the difficulty in studying these insects in the field, means that very little is known of their distribution and basic biology (Heads and Taylor 2012;Baena-Bejarano 2015).

Material and methods
The material studied here is deposited in the Instituto de Investigaciones Alexander von Humboldt, Villa de Leyva (IAvH-E) and the entomological museum of Universidad del Valle, Cali (MUSENUV). The male terminalia and the phallic complex were dissected and stored under glycerin in microvials mounted on the pin beneath the specimen. Some specimens were kept in alcohol. The description of morphological characters follows Heads (2010) and Heads and Taylor (2012) (Fig. 1). Interocular distance was measured using a calibrated micrometer slide adapted to a stereomicroscope. Other measurements were made from photographs analyzed with a calibrated digital scale in the program tpsdig2 (Rohlf 2006). Morphological characters were defined and documented in a Delta matrix and the description developed using Delta software (Dallwitz 1980(Dallwitz , 1999. Photographs were taken with a Leica digital camera attached to a stereomicroscope and focus-stacked in CombineZP (Hadley 2009). Scanning electron micrographs were produced using FEI Quanta 200 scanning electron microscope. Drawings were produced in Adobe Illustrator CS5 and Photoshop CS5. Paratypes. Five specimens from same locality as holotype: 1) ♂ (no. IAvH-E 137238), specimen preserved in alcohol; 2) ♂ (no. IAvH-E 137239), specimen preserved in alcohol; 3) ♀ (no. IAvH-E 137240), specimen preserved in alcohol; 4) ♀ (no. IAvH-E 137241), specimen preserved in alcohol; 5) ♀ (no. IAvH-E 142878), specimen dried and pinned. Specimens deposited at same institution as holotype.

Diagnosis.
The new species is almost cryptically similar to R. forceps Saussure, 1896 in that the uncus is elongate and strongly recurved, and that the median ridge of the male subgenital plate is produced distally, forming a short and densely setose bilobed process. However, it can be readily distinguished from the latter species by [1] antennae with white spots on flagellomeres 3-4 and 6-7 with flagellomere 5 entirely white; [2] epiproct produced laterally near its base; [3] brachium curved along its entire length without prominent apical bulge; and [4] phallic complex with virga thickened distally and not reaching beyond the membrane.
Head. Interocular distance more than half the eye width. Median ocellus fully developed. Patch that circumscribes anterodorsal margin of compound eyes absent. Internal margin of compound eyes convergent dorsally. Maxillary palp black, five segmented, with second segment reduced. Labial palp black. Gena below the compound eye and antennae insertion black.
Antennae black and filiform. Number of antennae segments 10. Scape wider than pedicel. Pedicel as long as 1 st flagellomere. White spot on scape absent. White spot on pedicel absent. White dorsal spot on flagellomere 1 and 2 absent. White dorsodistal spot on flagellomere 3 present. Flagellomere 4 white with a brownish slender anterior ring. Flagellomere 5 completely white. Flagellomere 6 white with a brownish slender distal ring extended ventrally to the segment half. Flagellomere 7 and 8 black.
Thorax. Pronotum black. Mesonotum black. Tegmina black. Hind wings with white, transverse groove. Procoxa black. Profemora black with an inner distal white spot. Protibiae black with three distal spines and an anterior external white rounded spot close to tibiae-femora articulation. Mesocoxa black. Mesotrochanter black. Mesofemora black. Mesotibiae black. Metafemora black. Semi-lunar process brown. Metatibia brown. Metatarsi brown and longer than metatibial posterior spurs.
Basal plate heavily sclerotized, long, basally strongly widened and distally strongly split. Cingulum with apodemes elongate and well-sclerotized. Sclerotized region of cingulum discontinuous with a distal membranous region in-between. Virga very slender near base and distally thickened. Virga not extended beyond cingulum (Fig. 2B).
Female. Body similar to male, except for antennal sexual dimorphism and abdominal sexual structures. White dorsodistal spot often present on flagellomere 2. Flagellomere 4 to 7 completely white. Subgenital plate smooth with two distal notches forming a rounded lobe in middle (Fig. 2C). The color is a lighter brown close to the notches.
Etymology. The specific epithet is patronymic and honours Señor Diego Baena, father of the senior author, in thanks for his care and dedication.
Distribution. This species is currently known from the type locality. Sympatric species. The new species was found in one of the malaise samples together with R. guacharoensis and R. ecuadoriensis, with which it is believed to live sympatrically.
Remarks. Ripipteryx diegoi sp. n. is assigned to the Forceps group based on the predominately black coloration, the form of the subgenital plate, morphology of the phallic complex and the body size 7.2-8.7 mm. This species is similar to R. forceps with which it shares the form of the subgenital plate presenting a median ridge forming a bilobed setose process in ventral view (Fig. 2G). This character allows differentiating it from the other species of the group. Moreover, it differs from R. forceps by the shape of the terminalia (Fig. 2D) where the epiproct is produced laterally near its base (Fig.  2E), the brachium in lateral view distally curved without prominent apical bulge; (see Günther 1969). Also, the new species differs in the color pattern of the male antennae with white spots on flagellomeres 3 to 7 of R. diegoi while these are present on the flagelomeres 1, 2, 4 and 5 of R. forceps.  Description. Male (holotype). Body length including wings 7.55 mm; pronotum length 1.38 mm, pronotum width 1.64 mm; tegmina length 2.91 mm; hind wings length 5.84 mm; interocular distance 0.37 mm. (n=1) (Fig. 3A).

Ripipteryx guacharoensis
Head. Interocular distance more than half the eye width. Median ocellus fully developed. Internal margins of compound eyes convergent dorsally. Slight yellowishwhite spot in the superior eyes corner. Gena below compound eye and antennae insertion black and below eye slightly yellowish. Maxillary palp black, distally slightly yellowish white, five segments with second reduced, fifth with strong setaes. Labial palp black.
Antennae thick and mainly black. Number of antennal segments 10. Scape wider than pedicel. Pedicel as long as 1 st flagellomere. Slight white distal spot on scape. White dorsal spot on pedicel. White dorsal spot on flagellomeres 1, 2 and 3. White dorsodistal spot on flagellomere 4. Flagellomeres 5, 6 and 7 black. Flagellomere 8 black with distal half completely white.
Thorax. Pronotum black with an anterior slender white line and an almost imperceptible yellowish at anterior corners. Tegmina black. Hind wing with white, transverse groove. Procoxa black. Protochanter black. Profemora black with a yellowish serrated distal inner lobe. Protibiae black with three distal spines. Mesocoxa black. Mesotrochanter black. Mesofemora black. Mesotibiae black distally brownish. Metafemora black; Semi-lunar process, metatibia and metatarsi brown.
Basal plate heavily sclerotized, very short and widened basally. Cingulum distally serrated without apodemes. Lateral valves pointed and serrated. Virga thick, distally rounded and serrated, basally with two slight tips. Ventral plate concave with a dorsal elevation in middle extended to virga (Fig. 3B).
Female. Body similar to male except for abdominal sexual structures. Subgenital plate obtuse (Fig. 3C).
Etymology. The specific epithet is derived from the name of the type locality, Parque Nacional Natural Cueva de los Guácharos.
Distribution. This species is currently known from the type locality. Sympatric species. This species was found in one of the malaise samples with the species R. diegoi and R. ecuadoriensis, which are believed to occur sympatrically.
Remarks. Ripipteryx guacharoensis sp. n. is assigned to the Marginipennis group based on the characters of the phallic complex, such as the very short and broad basal plate, the cingulum without apodemes, the presence of lateral valves, and the thickened virga (Fig. 3B).
The new species is similar to R. femorata in that both share a serrated distal inner lobe on the profemora, the shape of the male brachium in lateral view, the uncus reduced without distal hook and similar phallic complex (see Günther 1969). Nevertheless, it differs in the form of the ventral plate, which is concave in R. guacharoensis but is straight in R. femorata. The basal shape of the virga presents two slight tips in R. guacharoensis while in R. femorata it presents two strong and elongate tips (see Günther 1969). The most significant character separating both species is the posterior margin of the epiproct, which is triangular in R. guacharoensis (Fig. 3G) but parabolic in R. femorata (see Günther 1969).
According with Günther (1969) R. femorata is closely related to R. vicina and R. difformipes. Ripipteryx guacharoensis shares with these three species the form of the subgenital plate that in males is distally constricted with conspicuous long and curved bristles, supporting a probable relationship.  ing placement in the Crassicornis Group. It shares the presence of numerous sharp spiculae on the cingulum with R. antennata and R. atra and the antennal color pattern with R. atra. However, R. gorgonaensis differs from the former species by the absence of modifications of the antennae. In other members of the Crassicornis Group, certain antennomeres are fused (e.g. in R. atra) or otherwise modified (e.g. flattened and wide in R. laticornis and R. antennata); this is not the case in R. gorgonaensis. The latter is easily distinguished from other species of the Crassicornis and Forceps groups by the form of the terminalia (Figs 4C, D).

Ripipteryx gorgonaensis
A number of soft-bodied mites were found between the metanota and abdomina of some individuals. These are presumed to be ectoparasitic though further research is needed to clarify their biology and interaction with R. gorgonaensis (O. Combita pers. comm.).

Discussion
Five species groups had been proposed in the genus Ripipteryx which are largely defined by the morphology of the male terminalia and the phallic complex (Günther 1969;Heads 2010). Of the species described herein, Ripipteryx diegoi sp. n. and R. guacharoensis sp. n. can be confidently assigned to the Forceps and Marginipennis species groups respectively based on coloration, body size and the morphology of the male terminalia and internal genitalia. In contrast, the species group placement of R. gorgonaensis is problematic due to the presence of characters found in both the Crassicornis and Forceps groups such as modified subgenital plate and brachium, distal half of phallic complex weakly sclerotized, long apodemes of the cingulum, virga long and slender. Ripipteryx gorgonaensis was assigned to the Crassicornis group because it shares several characters of the terminalia with the species R. atra, R. antennata and R. laticornis and possesses spines on the cingulum like other species in the group. However, it lacks antennal modifications (a diagnostic character of the Crassicornis group) with the antennae more similar to those of Forceps group species. In briefly reviewing Günther's (1969) species group classification, Heads (2010) noted that the monophyly of some of the groups is questionable. Preliminary morphological phylogenetic analysis of the genus (Baena-Bejarano, unpublished) suggest that this is indeed the case, but more morphological and molecular data and a comprehensive phylogenetic treatment are required before a refined classification can be presented.
Montealegre, and anonymous reviewers for their valuable comments and critique of the manuscript, and Hugo Sánchez of the Universidad Nacional de Colombia for assistance with SEM. Partial financial support for this work was provided by Dirección de Investigación Sede Bogotá (DIB) grant 15221 entitled Relaciones filogenéticas de los géneros y grupos de especies de Ripipterygidae (Orthoptera: Tridactyloidea) basadas en morfología; and funding from the Instituto de Investigaciones de Recursos Biológicos Alexander von Humboldt (IAVH) as part of Gestión de Información y Conocimiento (GIC) program. Further thanks are due to Sina Firmenich for assisting the senior author with English translation, and Ranulfo González for his collecting effort.