Aclodes guane, Carvalho & Ferreira, 2025

Carvalho, Pedro Henrique Mendes & Ferreira, Rodrigo Lopes, 2025, Two new species of Aclodes Hebard, 1928 (Orthoptera: Phalangopsidae: Paragryllinae) from caves in El Peñón, Central-East Andean Colombia, Zootaxa 5590 (3), pp. 301-324 : 303-313

Aclodes guane sp. nov.

( Figures 1–6 , 7–15 , 16–19 , 20–22 , 23–29 , 30–31 , 32–36 ; Tables 1 and 2 )

Etymology —The specific epithet refers to the Guane people, a pre-Columbian South American indigenous group who used the Hoya de Casildo cave - one of the locals where the newly described species was found—as a ritualistic burial site.

Material examined — Holotype ( ISLA 126145 ), ♂, Colombia, Santander, El Peñón, Cueva de Los Carracos cave (73°49’42.88”W, 6°4’54.07”N), 17.viii.2023, R.L. Ferreira; condition: genitalia dissected, left hind-leg detached and store alongside the holotype . Allotype ( ISLA 126146 ), ♀, same data as the holotype; condition: right middle-leg missing . Paratypes: ( ISLA 126143 , ISLA 126144 , ISLA 126147 , ISLA 126148 , ISLA 126149 ), 2 ♂♂ and 3 ♀♀, same data as the holotype ; ( ISLA 126150 , ISLA 126151 , ISLA 126152 ), 1 ♂ and 2 ♀♀, Colombia, Santander, Bolívar, Hoya de Casildo cave (73°46’35.45”W, 6°4’49.17”N), 22.iii.2023, R.L. Ferreira .

Comments. Paratype’s condition varied: all male genitalia dissected, all male right tegmen detached and stored alongside the paratypes; ISLA 126143, ISLA 126151—left legs and right hind-leg detached and stored alongside the paratypes; ISLA 126144—left fore and middle-legs detached and stored alongside the paratype, left hind leg missing; ISLA 126147—intact; ISLA 126148—right hind-leg detached and stored alongside the paratype; ISLA 126149—left and right hind-legs detached and stored alongside the paratype; ISLA 126150—left and right hind-legs missing; ISLA 126152—right hind-leg missing.

Diagnosis —Combination of the following characteristics: posterior portion of the pseudepiphallic median lophi m-shaped; apical lobes of the pseudepiphallic median lophi curved dorsally, tapered at the apex, and ventrally covered in setae; posterior projection of the dorsal pseudepiphallic parameres as long as the posterior projection of the ventral parameres; ectophallic apodeme with a distinct apex that bends abruptly away from the center of the genitalia; ectophallic lateral bars sclerotized and sinuous, with a mall acute projection on the ventral inner margin near the apex, pointing towards the center of the genitalia; anterior portion of the endophallic sclerite elongated and V-shaped.

Male genitalia (holotype ISLA 126145, Figs 1–6 )—Phallic complex longer than broad and medially broadened. Pseudepiphallus: median lophi divided into anterior and posterior portion; anterior portion sclerotized, convex, with a secondary, less sclerotized dorsally projected bump in the center, anterior margin rounded, slightly projected and inclined towards the anterior region of the body ( Figs 2 and 3 , Ps.ap); posterior portion sclerotized, central region more sclerotized than the rest of the structure, lateral segments of the junction area with the median lophi anterior portion concave, posterior margin m-shaped, covered in small setae, and bearing two lateral projections— the pseudepiphallic apical lobes ( Figs 2, 4 and 6 , Ps.pp); apical lobes sclerotized, moderately elongated, sinuous in ventral view, curved dorsally and slightly away from the center of the genitalia, broadened at the base and tapered at the apex, which is fin-shaped (in lateral view) and ventrally covered in setae ( Figs 1 to 3, Ps.al); arms sclerotized, subtriangular, broadened laterally, and posteriorly connected to the pseudepiphallic apical lobes ( Figs 2 and 3 , Ps.arm ); ventral parameres sclerotized, exhibiting a dorsal projection running parallel to the dorsal parameres (Ps. p2), characterized by a rounded apex pointing towards the posterior end of the genitalia while inclined towards its center, ventrally subtriangular (in ventral view), articulated with the ectophallic lateral bars (Ect.lb), region near the apex of the ectophallic lateral bars internally concave and more sclerotized than the rest of the structure ( Figs 1, 3 and 6 , Ps.p1); dorsal parameres sclerotized, ventrally concave, inner margin more sclerotized when compared to the remainder of the structure, with a small and rounded ventral projection, posterior projection as long as the ventral paramere (Ps.p1) dorsal projection, with a rounded apex pointing towards the posterior end of the genitalia while inclined towards its center ( Figs 1 and 6 , Ps.p2); rami elongated, longer than the ectophallic apodeme (Ect. ap), broadened at the base, tapered at the end, S-shaped in lateral view, proximal half slightly externally concave, distal half internally concave, apex curving towards the central axis of the genitalia ( Figs 1 to 3, R). Ectophallic invagination: ectophallic arc well-developed, sclerotized, dorsally projected, central portion ventrally concave, located between the pseudepiphallic posterior portion and the base of the endophallic sclerite, posterior margin connected to the membranous sheath surrounding the endophallic sclerite posterior portion (End.sc.p), anterior margin irregular, forming an arc characterized by a broad and quadrangular central retraction ( Fig. 2 , Ect.arc); apodemes slightly sclerotized, elongated, slightly longer than the endophallic sclerite, inclined towards the center of the genitalia, apex curving away in the opposite direction (in dorsal view) ( Figs 2 and 3 , Ect.ap); lateral bars well-developed, sinuous, sclerotized, ventrally projected, reaching the ventral apex of the ventral paramere, broadened near the apex, which is fin-shaped, ventral inner margin with a small acute projection towards the center of the genitalia ( Figs 1 to 5, Ect.lb). Endophallus sclerite: anterior portion well-developed, sclerotized, V-shaped (in ventral view), with a long membranous apodeme on the opposite side of the central groove, akin to a keel ( Figs 1 and 6 , End.sc.a); duct short and highly sclerotized, its base roughly heart-shaped in ventral view ( Figs 1 and 6 , End. sc.d); posterior portion also heart-shaped, inserted in the central ventral concavity of the pseudepiphallic median lophi posterior portion, but not connected to it, surrounded by a trapezoidal and membranous sheath, which has in its posterior central portion a U-shaped sclerotized region ( Figs 1, 5 and 6 , End.sc.p).

Variations in male genitalia (paratypes ISLA 126143, ISLA 126144, ISLA 126150)—Pseudepiphallic dorsal (Ps.p2) and ventral (Ps.p1) parameres exhibit slight variations in their inclination relative to the longitudinal axis of the phallic complex; ectophallic lateral bars and pseudepiphallic ventral parameres show slight variations in the angle formed by the two contacting structures (in lateral view); apices of the ectophallic apodemes may be positioned closer together at the center of the genitalia or farther apart, near the internal face of the pseudepiphallic rami; apices of the ectophallic apodemes can be rounded, as in the holotype, or acute; the position of the endophallus (and consequently, the ectophallic arch that accompanies it) varies, being either interiorized or exteriorized, projecting slightly beyond the pseudepiphallic median lophi posterior portion, as in the holotype, or retracted between the parameres of the pseudepiphallus; posterior portion of the endophallus may be less sclerotized, semicircular in shape and exhibit two small, sclerotized, ear-like folds on each side (similar to those observed in Aclodes oscari sp. nov., but even more developed).

Male body morphology and color (holotype ISLA 126145, Figs 7–15 )— Body color: vertex brown, with four parallel anteroposterior yellowish-brown stripes; ocellar regions yellow; fastigium, frons, and subgena dark brown, covered with yellowish spots; dorsal portion of the gena is brown, the middle portion is lighter and yellowish-brown, while the ventral portion is dark brown, following the coloration pattern of the subgena; clypeus and labrum yellow, apex of labrum white; mandibles and laciniae yellowish-brown, with sclerotized apex; galeae, labium, maxillary palps, and labial palps yellowish-white; scape, pedicel, and flagellum brown; pronotum brown, divided in the middle by a yellowish-brown anteroposterior stripe and covered with yellowish-brown spots, with the lateral portion darker and the apex of lateral lobes light yellow; right tegmen yellowish-brown; tergites brown or dark brown, sprinkled with yellowish-brown and yellowish-white spots; sternites yellowish-white; supra-anal plate brown; subgenital plate dark brown, with a yellowish-white spot in the center and two oval black spots near the base; cerci yellowish-brown; legs exhibiting alternating dark and light rings, with the external face of the hind femur covered with brown stripes; tympanum white. Head ( Figs 8, 9 and 11 ): elongated dorsoventrally, slightly pubescent; fastigium covered with elongated setae and measuring approximately half the length of the antennal scape; maxillary palps pubescent, with palpomeres I and II short and globose, palpomeres III and IV elongated and of similar length, and palpomere V longer than the others with a clavate apex; labial palps also pubescent, with palpomeres I, II, and III progressively increasing in size, apex of palpomere III dilated; ocelli depigmented; compound eye slightly reduced, with a small portion of the dorsal region near the coronal suture depigmented. Thorax ( Figs 7 and 12 ): pronotum slightly pubescent, broader than long; anterior and posterior margins arched and covered with long setae; lateral lobes subtriangular, directed ventrally and slightly towards the posterior region of the body; metanotal glands absent. Abdomen ( Figs 12 to 15): pubescent; supra-anal plate approximately trapezoidal, slightly longer than the paraprocts, with a rounded posterior margin covered with long setae and lateral margins featuring a small invagination in the centroposterior portion; subgenital plate elongated, approximately twice as long as the supra-anal plate, rectangular in shape, with a slightly curved posterior margin, elongated setae, and a small central dent; cerci pubescent, with elongated setae throughout their length, and globose setae on the inner portion of their bases. Right tegmen ( Figs 10 and 12 ): slightly sclerotized, roughly oval, reaching the anterior portion of the fourth tergite; mirror almost rhomboid, with a complete crossvein perpendicular to the longitudinal axis of the tegmen and an incomplete crossvein near the posterior margin; harp with one incomplete and three complete crossveins; basal field 1A and 2A veins well-defined, converging and then dividing again into two distinct veins; lateral field with two parallel longitudinal veins and numerous weakly marked, anastomosed crossveins. Fore-legs (similar to Figs 16 and 17 — paratype, ISLA 126143): pubescent, proximal portion of the internal face of the tibia bearing an oval tympanum; apex with two ventral spurs; tarsomere serrated ventrally. Middle-legs (similar to Figs 16 and 17 — paratype, ISLA 126143): also pubescent, apex of the tibia with three spurs (two ventral and one lateral on the external face); tarsomere also serrated ventrally. Hind-legs (similar to Figs 18 and 19 — paratype, ISLA 126143): pubescent, femur developed; tibia longer than the femur, bearing two longitudinal rows of dorsal spines (left tibia with three external and four internal dorsal spines, right tibia with four dorsal spines on both sides), and two sets of apical spurs, three internal and three external (on the inner side, the most ventral spur, spur d, being extremely reduced); tarsomere I longer than the sum of the length of tarsomeres II and III, with one longitudinal, externally positioned, dorsal row of a few (four or five) short spines, plus one small spine on the apex of the inner dorsal side, and two apical spurs, with the internal spur being longer than the external one.

Variations in body morphology (paratypes)—Harp with five complete crossveins, stridulatory file with 109 teeth ( ISLA 126143, Fig. 20 ). Harp with one incomplete and four complete crossveins, stridulatory file with 99 teeth ( ISLA 126144, Fig. 21 ). Harp with five complete crossveins, stridulatory file with 117 teeth ( ISLA 126150, Fig. 22 ). Left and right tibiae III featuring two longitudinal rows of four dorsal spines ( ISLA 126143, ISLA 126144); tibia III outer row bearing five spines, the two most proximal close to each other ( ISLA 126147). Tympanum droplet-shaped ( ISLA 126143, Fig. 17 ). Supra-anal plate as long as the paraprocts ( ISLA 126143, ISLA 126144, ISLA 126150).

Female body morphology and color (paratype ISLA 126151, Figs 23–29 )—Slightly larger than the male. Body color: coloration pattern similar to the holotype, except for the vertex, pronotum, and supra-anal and subgenital plates; vertex with two yellowish-brown anteroposterior stripes converging at the base of the coronal suture, where there is a rounded yellowish spot; pronotum with two yellowish-brown oval spots in the posterior portion; supra-anal plate brown, with three well-defined regions separated by a central whitish ring ( Fig. 25 ); subgenital plate yellowish-white ( Fig. 26 ); ovipositor orange-brown, with a dark brown apex. Tegmina ( Fig. 24 ): reduced, without apparent veins, rounded, positioned on the sides of the mesonotum, reaching the anterior margin of the metanotum. Legs ( Fig. 23 ): similar to those of the holotype; proximal portion of the internal face of tibia I bearing an oval tympanum; dorsal face of tibia III with two rows of four spines (outer row of right tibia III lacking the most proximal spine, its insertion point present); sets of apical spurs equal to those of the holotype. Ovipositor ( Figs 27 to 29): elongated, longer than the cerci, lance-shaped, slightly curved ventrally, with a sclerotized, tapered apex, triangular in dorsal view. Copulatory papilla: membranous.

Ecological remarks —Los Carracos Cave is situated in a semi-preserved karst area, featuring patches of forested regions surrounded by pastures ( Fig. 32 ). This limestone cave extends over 1.5 kilometers of mapped galleries, comprising two levels: one partially active (trespassed by a drainage) and the other inactive. The cave has two entrances accessing the upper level ( Fig. 33 ), located very close to each other, that lead to a 15-meter-wide gallery descending to a lower entrance. A small stream flows from a waterfall formed by epikarst infiltration ( Fig. 35 ) between the entrances, partially disappearing into a lateral gallery. In the anterior part of this first gallery, a low opening provides access to a 10-meter pit, which in turn leads to the lower level. As wide as the upper gallery but more horizontal, this lower section extends upstream for several hundred meters until reaching a calcitic fill. The upper level of the cave features highly ornamented areas with numerous speleothems, making it a significant tourist attraction in the region ( Fig. 34 ).

While the upper level of the cave is oligotrophic, the lower level hosts a large colony of oil birds ( Steatornis caripensis ), which produce a significant amount of guano. During our sampling in the cave, we did not access the lower level, so all observed crickets were found on the upper level. However, given the abundance of organic resources in the lower level, it is likely that the crickets also inhabit that part of the cave.

Specimens of Aclodes guane sp. nov. were frequently observed on the cave floor and walls of the upper level ( Figs 30 and 31 ). However, they were sparsely distributed, and specimens were not commonly found near each other. It is important to note that additional specimens of Aclodes guane sp. nov. were discovered in another cave, Hoya de Casildo cave, located approximately 5.7 km away in a straight line from Carracos cave. Despite being much smaller than Carracos cave, Hoya de Casildo cave is significant as an ancient ritualistic site used by the Guane people for burying their dead. Along the cave, particularly in the last lower chamber, where the crickets were most abundant, there are several human remains, some of which are partially calcified, dating back more than 2,000 years ( Fig. 36 ).

It is important to note that we only visited a few caves during our expedition to the El Peñon area. Considering the distance between Los Carracos and Hoya de Casildo caves, it is plausible to assume that other populations of Aclodes guane sp. nov. might exist. Lastly, similar to La Tronera cave, human activities such as deforestation for pastureland, limestone extraction, and unregulated tourism inside the caves may impact the caves and consequently their fauna.

Figures

FIGURES 1–6. Aclodes guane sp. nov. (holotype, ISLA 126145) male genitalia. (1) ventral view, (2) dorsal view, (3) lateral view, (4) anterolateral view, (5) posterolateral view, (6) frontal view. Abbreviations: Ps.ap, pseudepiphallic anterior portion; Ps.pp, pseudepiphallic posterior portion; Ps.al, pseudepiphallic apical lobe; Ps.arm, pseudepiphallic arm; Ps.ml, pseudepiphallic median lophi; Ps.p1, pseudepiphallic ventral paramere; Ps.p2, pseudepiphallic dorsal paramere; R, pseudepiphallic rami; Ect. ap, ectophallic apodeme; Ect.arc, ectophallic arc; Ect.lb, ectophallic lateral bar; End.sc.a, endophallic sclerite anterior portion; End.sc.d, endophallic sclerite duct; End.sc.p, endophallic sclerite posterilor portion.

FIGURES 7–15. Aclodes guane sp. nov. (holotype, ISLA 126145) body morphology. (7) pronotum in dorsal view; (8) head in dorsal view; (9) head in lateral view; (10) tegmina in dorsal view, mirror highlighted in green, harp highlighted in blue, stridulatory file, 1A and 2A veins highlighted in pink; (11) head in frontal view; (12) pronotum and tegmina in lateral view; (13) abdomen apex in lateral view; (14) supra-anal plate in dorsal view; (15) subgenital plate in ventral view.

FIGURES 16–19. Aclodes guane sp. nov. (paratype, ISLA 126143) left legs. (16) legs I and II, outer view, apical spines of tibia II highlighted; (17) legs I and II, inner view, tympanum of tibia I and apical spine of tibia II highlighted; (18) leg III, inner view, apical spurs of the tibia highlighted; (19) leg III, outer view, apical spurs of the tibia highlighted.

FIGURES 20–22. Aclodes guane sp. nov. (paratypes) right tegmina. (20) ISLA 126143, dorsal and lateral views; (21) ISLA 126144, dorsal and lateral views; (22) ISLA 126150, dorsal and lateral views.

FIGURES 23–29. Aclodes guane sp. nov. (paratype, ISLA 126151) female body morphology. (23) tympanum, tibia I, inner view; (24) pronotum and tegmina, dorsal view; (25) supra-anal plate in dorsal view; (26) subgenital plate in ventral view; (27) ovipositor in lateral view; (28) ovipositor apex in lateral view; (29) ovipositor apex in dorsal view.

FIGURES 30–31. Aclodes guane sp. nov. (30) adult male, Los Carracos cave; (31) adult female, Los Carracos cave.

FIGURES 32–36. Aclodes guane sp. nov. habitat. (32) Los Carracos cave external environment; (33) Los Carracos cave entrance; (34) Los Carracos cave internal environment, speleothems; (35) Los Carracos cave internal environment, small stream formed by epikarst infiltration; (36) Hoya de Casildo cave, human remains partially calcified, Guane burial site.