Research Article |
Corresponding author: Atilano Contreras-Ramos ( acontreras@ib.unam.mx ) Academic editor: Davide Badano
© 2023 Adrian Ardila-Camacho, Shaun L. Winterton, Atilano Contreras-Ramos.
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.
Citation:
Ardila-Camacho A, Winterton SL, Contreras-Ramos A (2023) The genus Climaciella Enderlein, 1910 (Neuroptera, Mantispidae) in French Guiana. ZooKeys 1153: 37-64. https://doi.org/10.3897/zookeys.1153.95960
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The genus Climaciella Enderlein, 1910 is a remarkable group of mantidflies (Neuroptera: Mantispidae: Mantispinae) distributed from Canada to Argentina, including parts of the Caribbean. This genus comprises nine valid extant species plus an extinct species from the late Oligocene of France. Species exhibit Batesian mimicry with vespid wasps (Vespidae). Herein, six species of Climaciella from French Guiana are documented. Before this study only C. semihyalina (Le Peletier de Saint Fargeau & Audinet-Serville in
Lacewings, Neotropics, taxonomy
Mantispidae (mantidflies) are a family of predatory insects included within the superfamily Mantispoidea (
Among the raptorial Mantispoidea, Mantispinae is the most diverse and widely distributed subfamily (
The biology of Climaciella is known mostly from studies performed with C. brunnea (Say, 1824), a widely distributed species composed of a complex of morphs (or subspecies), which mimic polistine wasps and occur through North America, Mexico, and Central America (
After hatching, the first instar larva of C. brunnea adopts a vertical posture with the aid of the suctorial eversible process at the end of the abdomen while it maintains its legs extended (
The taxonomy of the genus Climaciella was addressed in the classical studies of Mantispidae by
Specimens studied herein were collected in different localities of French Guiana between 2014 and 2016 using light traps, and then deposited in California State Collection of Arthropods, Sacramento CA (
Subfamily Mantispinae Leach, 1815
Climaciella is distinguished from other mantispines of the New World by the antennal flagellomeres ≥ 3 times as wide as long in frontal view, the lateral parapsidal furrow not developed, and the female gonocoxites and gonapophyses VIII separated by a membranous strip. Additionally, the membrane of the anterior half of the wings is amber colored, and all the species exhibit Batesian mimicry with polistine wasps.
Climaciella amapaensis Penny, 1982, C. brunnea (Say, 1824), C. cubana Enderlein, 1910, C. elektroptera sp. nov., C. henrotayi Nel, 1989, C. nigriflava sp. nov., C. obtusa Hoffman, 2002, C. personata (Stitz, 1913), C. porosa Hoffman, 2002, C. rafaeli Calle et al. in Ardila et al. 2018, C. risaraldensis sp. nov., C. semihyalina (Le Peletier de Saint Fargeau & Audinet-Serville in
1 | Prothorax straight or at least only slightly bent medially in lateral view | 2 |
– | Prothorax bent medially in lateral view | 3 |
2 | Body entirely black | Climaciella sp. |
– | Body yellow with black stripes | Climaciella amapaensis Penny, 1982 |
3 | Head and prothorax black; anterior half of the wings smoky or dark amber | 4 |
– | Head and prothorax mostly yellow or entirely dark brown; wings with anterior half pale amber | 5 |
4 | Wings with anterior half smoky, the posterodistal margin of the pigmented area darker; abdomen orange | Climaciella tincta (Navás, 1914) |
– | Wings with anterior half dark amber and hyaline posterodistal, triangular area; abdomen black | Climaciella semihyalina (Le Peletier de Saint-Fargeau & Audinet-Serville, 1825) |
5 | Body mostly yellow with dark brown stripes and marks | Climaciella nigriflava sp. nov. |
– | Body entirely dark brown | Climaciella elektroptera sp. nov. |
French Guiana: Roura, Montagne des Chevaux, Carrière du Galion, Créte avec foret sur quartzite érodée, 04°44'31.54"N, 52°25'53.02"W, 16.V.2015, automatic light trap (blue) (1♀ CNIN).
Despite this species exhibits a similar coloration pattern as C. semihyalina, the straighter prothorax and the wing pattern easily differentiate the unknown species. Compared to C. semihyalina, the anterior half of both wings of the specimen examined have paler areas inside the cells and wing apexes (Fig.
French Guiana.
Climaciella amapaensis Penny, 1982: 450. Holotype: male, Brazil (DZUP).
French Guiana: Maripasoula, Mitaraka, Contreforts du Mitaraka, crique Alama, Foret vallonnée au pied d’inselbergs, 20.VIII.2015, automatic light trap (GemLight) (1 sex? CNIN).
Records of Climaciella amapaensis are scarce, but this species can be easily recognized by its coloration pattern that consists of a yellow body with black stripes and marks. Even though a similar color pattern is expressed by C. nigriflava sp. nov., C. cubana, C. risaraldensis sp. nov., and some morphs of C. brunnea, this species is distinguished because on the head there are two transversal bands at the level of the antennal insertion and on the upper portion of the region of the vertex, the compound eyes are enlarged and the region of the gena is narrow. In addition, the prothorax is straight in lateral view with longitudinal, lateral dark brown bands. This species is also unique within the genus due to the presence of narrow forefemur with thin processes. On the male genitalia, the apex of the gonocoxites X is not forked and blade-shaped, and the hypomeres are present as a single granule-shaped sclerite on the gonostyli X membrane laterally on each side.
This species is herein reported from French Guiana for the first time. In the original description, this species was reported from Brazil (Amapá) (
Brazil, French Guiana, Peru.
Holotype
♂, French Guiana: Maripasoula/Camopi, Mont Itupé, Massif tabulaire, Pente oust (600 m), 28.XI.2014, light trap (
This species is easily differentiated from its congeners based on its uniformly dark brown body coloration. The prothorax is bent medially like in C. semihyalina and C. obtusa and lacks a prominent hump as in C. porosa. The wing coloration pattern is similar to that of C. porosa, with the anterior half of both wings pale amber. On the male genitalia, the gonocoxites X are bifid at the apex, and the gonostyli X membrane is set with a granule-shaped hypomere laterally at each side. Additionally, the gonostyli X are ribbon-shaped and the female spermatheca is relatively short and simple.
Measurements. Head width: 2.9‒3.2 mm; Head length: 2.7‒3.4 mm; Prothorax length: 3.4‒3.9 mm; Forefemur length: 4.1‒6.4 mm; Forefemur maximum width: 1.7‒2.1 mm; Forewing length: 15.4‒16.5‒18.8 mm; Forewing maximum width: 3.5‒4.1 mm; Hindwing length: 13.3‒15.8 mm; Hindwing maximum width: 3.2‒3.9 mm.
Coloration
(Figs
Morphology
(Figs
Male terminalia
(Fig.
Female terminalia
(Fig.
The specific name of this species is from the Greek ἤλεκτρον (ḗlektron) meaning amber, and πτερόν (pteron), meaning wing, in allusion to the coloration of the wings of this species.
This new species is similar to C. porosa based on its general coloration; however, the bent prothorax and the lack of a hump on the medial region of the pronotum quickly differentiate it. Another species similar to C. elektroptera sp. nov. is C. personata from Bolivia, although the latter has a more patterned body coloration, and the posterior limit of the amber anterior area of the wings is darker and noticeably marked. Based on the bent prothorax, C. elektroptera sp. nov. is probably closely related to C. obtusa, C. semihyalina, C. personata, C. rafaeli, and C. nigriflava sp. nov.
Based on the coloration pattern, size, and distribution of C. elektroptera sp. nov., the possible wasp models for this species could be Agelaia angulata (Fabricius, 1804) or Polistes deceptor Schulz, 1905 (R. Lopes, pers. comm. 2022)
French Guiana.
Holotype
♂, French Guiana: Roura, Montagne des Chevaux, Carrière du Galion, Crete avec foret sur quartzite érodée, 04°44'31.54"N, 52°25'53.02"W, 22.II.2016, automatic light trap (blue) (
This new species has a similar body coloration pattern as C. amapaensis, C. cubana, and certain morphs of C. brunnea. However, the bent prothorax at midlength in lateral view distinguishes the new species from the aforementioned ones. Furthermore, the head in C. nigriflava sp. nov. is basically entirely yellow, while the remaining species with similar coloration exhibit dark stripes or marks. The coloration of the femur is also similar to that of C. amapaensis, yet the more robust femur with thickened processes distinguishes C. nigriflava sp. nov. from the former. Moreover, the wing coloration of this species is similar to that of C. porosa and C. elektroptera sp. nov. In the male genitalia, the gonocoxites X have the posterior apex truncate, the hypomeres are absent, and the gonostyli X are elongated, ribbon-shaped with the apex forming an obtuse angle.
Measurements. Head width: 3.0‒3.1 mm; Head length: 2.6‒3.0 mm; Prothorax length: 3.6‒3.8 mm; Forefemur length: 5.2 mm; Forefemur maximum width: 1.9 mm; Forewing length: 15.9‒16.5 mm; Forewing maximum width: 3.5‒3.7 mm; Hindwing length: 13.5‒14.1 mm; Hindwing maximum width: 3.2‒3.5 mm.
Coloration
(Figs
Morphology
(Figs
Male terminalia
(Fig.
Female terminalia
(Fig.
The specific epithet of this species is a combination of the Latin words niger which means black, and flavus meaning yellow, in allusion to the coloration pattern of the body of this species.
Based on the prothorax morphology, this new species appears to be closely related to C. obtusa, C. semihyalina, C. personata, C. rafaeli, and C. elektroptera sp. nov. The body coloration pattern of the new species with head and thorax predominantly yellow, and a large dark area on the abdomen indicates this species could be a mimic of the vespid wasps Agelaia pallipes (Olivier, 1792) or Mischocyttarus cerberus Ducke, 1918 in the case of smaller specimens, or Polistes testaceicolor Bequard, 1937 or Agelaia testacea (Fabricius, 1804) in the case of larger specimens (R. Lopes, pers. comm. 2022).
Mantispa semihyalina
Le Peletier de Saint-Fargeau & Audinet-Serville in
Mantispa chalybea
Erichson, 1839: 160, sex not indicated. Syntypes: sex unknown, Brazil, Suriname (
Mantispa grandis; Burmeister, 1839: 967. Not available with Burmeister as author. Synonymized by Westwood 1852: 253.
Nobrega tinctus
Navás, 1914: 233, sex not indicated (see C. tinctus below). Holotype: sex unknown, Brazil (NHMUK). Synonymized by
French Guiana: Maripasoula, Mitaraka, Contreforts du Mitaraka, crique Alama, Foret vallonnée au pied d’inselbergs, 12.III.2015, light trap (1♂ CNIN); same data but automatic light trap (blue) (1♂
The overall black coloration of the body of this species is shared with C. obtusa, C. rafaeli, and the synoeca morph of C. brunnea. However, the wing pattern of C. semihyalina easily separates it from the former two, whereas the bent prothorax distinguishes this species from the latter. This species is rapidly recognized by the dark amber anterior half of the wings, while in the posterior half, the basal region is pale amber, while a preapical triangular area is hyaline. On the male genitalia, the apex of the gonocoxites X is bifid, and the gonostyli X are spine-shaped.
Climaciella semihyalina, is the most commonly encountered species of the genus in South America, being widely distributed in the Amazon. The bent prothorax together the robust forefemora of this species resemble the shape of the head of its potential models, i.e., the vespids Polybia simillima Smith, 1862, P. ignobilis (Haliday, 1836) (
Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Paraguay, Peru, Suriname, Uruguay.
Nobrega tinctus Navás, 1914: 233. Holotype: sex unknown, Brazil (NHMUK).
Climaciella duckei Navás, 1915: 196. Holotype: sex unknown, Peru (NMBS). Synonymized by Alvim, 2021: 31.
Climaciella tincta
(Navás, 1914);
French Guiana: Maripassoula/Camopi, Mont Itoupé, Massif tabulaire, Pente oust (600 m), 23.XI.2014, light trap (1♂, 1♀ CNIN); same data but 24.XI.2014, light trap (1♀
This species has a distinctive coloration of the body with black head and thorax and orange abdomen. The prothorax is bent like in C. semihyalina, C. obtusa, C. personata, C. rafaeli, and the new species described herein. The wings have smoky anterior half, with the posterodistal margin of this pigmented area more marked. On the male genitalia, the apex of the gonocoxites X is bifid, the hypomeres are present as a lateral granule-shaped sclerite at each side, and the gonostyli X is elongated and narrow.
Based on the prothorax morphology, C. tincta is apparently closely related with C. semihyalina, C. obtusa, C. personata, C. rafaeli, C. elektroptera sp. nov., and C. nigriflava sp. nov. This species was originally described by
Based on the distribution and coloration pattern exhibited by C. tincta (Fig.
Brazil, French Guiana.
Holotype ♂, Colombia: Risaralda, Tatamá, Centro de visitantes planes de San Rafael, 5°4'20,87"N, 75°57'44,45"W, 2400 m, 10.VI.2010, F. Gaviria, entomological net (CEH-085).
This new species may be distinguished from the majority of the species in the genus by the general coloration pattern that is yellow with black stripes and marks. Furthermore, C. risaraldensis sp. nov. is easily separated from C. nigriflava sp. nov. by the black vertex region and the straight prothorax. On the other hand, this new species can be separated from C. amapaensis by having smaller compound eyes and broadened region of the gena, pronotum dark brown with small diffuse yellow areas, anterior half of the wing pale amber, the forked apex of the male gonocoxites X, and hypomeres present as two granules on each side of the gonostyli X membrane.
See
This species is named after the department of Risaralda, a region of the central Andes of Colombia where the type locality of this species is located.
Based on the descriptions of
Based on the morphology of the prothorax, this species is probably closely related to C. amapaensis, C. brunnea, C. cubana, and C. porosa. Among these, the new species is similar to certain morphs of C. brunnea; however the wing coloration, the male pregenital apparatus, and the genitalic morphology are markedly differentiated between both species.
Colombia.
In the present study, the number of species of Climaciella in French Guiana is increased from one to six, based on considerable collecting efforts performed in this small portion of the Amazon. The diversity of species in this territory is quite remarkable compared to other, larger Neotropical countries. In Colombia and Brazil for example, five species of this genus have been reported to date (
The two Guianese new species described herein are apparently closely related based on genitalic morphology. Both species appear to form a group of species together with C. semihyalina, C. obtusa, C. rafaeli, C. personata, and C. tincta based on the bent prothorax, a hypothesis supported in the cladistic analysis performed by
Climaciella was recovered as sister of the remainder of mantispine genera of the New World by
We thank Rogério Botion Lopes for his help recognizing the possible wasp models of the species studied herein. Sincere thanks to Renato Machado for sharing information about the genus Climaciella from Brazil, to Michael Ohl for kindly providing high resolution images of the holotypes of C. personata and C. tincta, as well as for useful reviews to this contribution, and to associate editor D. Badano for encouraging feedback on a previous manuscript version. AAC acknowledges Posgrado en Ciencias Biológicas, UNAM, for support during his doctoral studies, and Consejo Nacional de Ciencia y Tecnología (CONACyT-Mexico) for support through a doctoral scholarship. We thank Susana Guzmán-Gómez (Laboratorio de Microscopía y Fotografía de la Biodiversidad, LaNaBio-IBUNAM) for help with high resolution stereomicroscope photography. AAC and ACR acknowledge Instituto de Biología, UNAM for general support for this research. General support is also acknowledged through the project “Biodiversidad de Neuroptera en México: un enfoque taxonómico integrativo” (CONACYT CB2017-2018, A1-S-32693) and “Biodiversidad de grupos selectos de Neuropteroidea de la Península de Baja California” (PAPIIT IN209721). Specimens were collected under ABS permit number APA 973-1 as part of the “Our Planet Reviewed” Guyane-2015 expedition in the Mitaraka range, in the core area of the French Guiana Amazonian Park, organized by the