Research Article |
Corresponding author: Ivanklin S. Campos-Filho ( ivanklin.filho@gmail.com ) Academic editor: Matthew L. Niemiller
© 2022 Ivanklin S. Campos-Filho, Jéssica S. Gallo, Jonas E. Gallão, Dayana F. Torres, Lília Horta, Yesenia M. Carpio-Díaz, Carlos M. López-Orozco, Ricardo Borja-Arrieta, José O. Aguiar, Maria E. Bichuette.
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:
Campos-Filho IS, Gallo JS, Gallão JE, Torres DF, Horta L, Carpio-Díaz YM, López-Orozco CM, Borja-Arrieta R, Aguiar JO, Bichuette ME (2022) Unique and fragile diversity emerges from Brazilian caves – two new amphibious species of Xangoniscus Campos-Filho, Araujo & Taiti, 2014 (Oniscidea, Styloniscidae) from Serra do Ramalho karst area, state of Bahia, Brazil. Subterranean Biology 42: 1-22. https://doi.org/10.3897/subtbiol.42.75725
|
Two new troglobitic species of Xangoniscus are described from two caves of Serra do Ramalho karst area, Bambuí geomorphological group, state of Bahia. Xangoniscus lapaensis sp. nov. is described from Gruna Boca da Lapa cave, and X. loboi sp. nov. from Gruna da Pingueira II cave. Both species are blind and depigmented and show amphibious habits, as observed for all species of Xangoniscus described until now. Xangoniscus lapaensis sp. nov. occurs in travertine pools fed by water of the upper aquifer, and X. loboi sp. nov. occurs in a small stream, an upper vadose tributary. Both species occur in fragile microhabitats. Ecological and behavioral data, conservation remarks, and IUCN conservation assessments are included to provide background data for conservation efforts in this unique karst area.
Bambuí geomorphological group, Limestone, Terrestrial isopods, Troglobitic, Upper Aquifer
Terrestrial isopods (Oniscidea) are the only crustacean lineage completely adapted to the terrestrial way of life (
Terrestrial isopods are one of the most representative taxa in the Brazilian subterranean environments, probably due to the special environmental conditions, such as high humidity, substrate heterogeneity, and absence of specialized predators (
Among the troglobitic representatives of the Brazilian Oniscidea, 12 species have an amphibious way of life, all in the Styloniscidae: Chaimowiczia uai Cardoso, Bastos-Pereira, Souza & Ferreira, 2021, C. tatus Cardoso, Bastos-Pereira, Souza & Ferreira, 2021, Iuiuniscus iuiuensis Souza, Ferreira & Senna, 2015, Spelunconiscus castroi Campos-Filho, Araujo & Taiti, 2014, Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014, X. ceci Cardoso, Bastos-Pereira, Souza & Ferreira, 2020, X. dagua Cardoso, Bastos-Pereira, Souza & Ferreira, 2020, X. ibiracatuensis Cardoso, Bastos-Pereira, Souza & Ferreira, 2020, X. itacarambiensis Bastos-Pereira, Souza & Ferreira, 2017, X. lundi Cardoso, Bastos-Pereira, Souza & Ferreira, 2020, X. odara Campos-Filho, Bichuette & Taiti, 2016, and X. santinhoi Cardoso, Bastos-Pereira, Souza & Ferreira, 2020.
The present work describes two new species of Xangoniscus from the Serra do Ramalho karst area, state of Bahia, both occurring in special microhabitats, namely the upper aquifers fed by infiltration of water in the limestone rock. Moreover, aspects of their ecology and natural history, as well as conservation remarks about this particular and unique fauna and the region of Serra do Ramalho are given.
Specimens of Xangoniscus were collected by hand with the aid of tweezers and brushes. Specimens were stored in 70% and 100% ethanol. Descriptions are based on morphological characters with the use of micropreparations in Hoyer’s medium (
Maps were produced highlighting the caves where both Xangoniscus species occur, as well the hydrological attributes and pressures of economic activities in the region, representing impacts to the cave fauna. The distribution map was constructed with QGIS software (v. 3.18.1) and was edited in PowerPoint Microsoft 365 (v. 2108).
The material is deposited in the scientific collection of cave fauna of the Laboratório de Estudos Subterrâneos (LES), Universidade Federal de São Carlos, São Carlos, Brazil (Curator: Maria E. Bichuette).
Both populations were studied in one field trip in October 2020, during the dry-wet season transition. Specimens were sampled ad libitum (sensu
To assess population parameters (density and size of the population), the individuals were counted along microhabitat inside caves, and measurements (such as length, width, and total area) of the occurrence sites were taken. The population parameters were calculated by dividing the number of individuals observed by area of the occurrence site. Photographs and short videos were taken to document the behavior of both species and the influence of the microhabitat on their distribution in the caves (see Supplementary Online Material).
The Serra do Ramalho karst area, southwestern State of Bahia, is located within the Bambuí geomorphological group, which hosts a high number of caves, mostly without legal protection. Some caves within this karst region extend for more than 15 km, developing huge cave systems (
Maps of Serra do Ramalho karst area, State of Bahia, northeastern Brazil A limestone outcrops of the Bambuí geomorphological group, Fraturado and Urucuia-Areado aquifer systems showing the type-localities (caves) of X. lapaensis sp. nov. and X. loboi sp. nov. B headwaters and São Francisco river basin C pressure for land use showing human-caused alterations D pressure for mining activities.
Natural habitat and habitus of Xangoniscus lapaensis sp. nov. A entrance of Gruna Boca da Lapa cave B travertine pools where Xangoniscus lapaensis sp. nov. occurs C specimen of Xangoniscus lapaensis sp. nov. submerged on rocky substrate at the travertine pools D specimens of Xangoniscus lapaensis sp. nov. submerged on silt and rocky substrate.
Natural habitat and habitus of Xangoniscus loboi sp. nov. A karst area where Gruna da Pingueira II cave is located B karst and entrance of Gruna da Pingueira II cave C specimen of Xangoniscus loboi sp. nov. submerged on the silt substrate D damn inside the cave E specimens of Xangoniscus loboi sp. nov. feeding on seed fruits F specimens of Xangoniscus loboi sp. nov. submerged on silt and rocky substrate G specimens of Xangoniscus loboi sp. nov. submerged on the rocky substrate H stream inside Gruna da Pingueira II cave where Xangoniscus loboi sp. nov. occurs.
Xangoniscus aganju Campos-Filho, Araujo & Taiti, 2014, by original designation.
Bahia, Feira da Mata, Gruna Boca da Lapa cave (13°56'46.4"S, 44°11'12.0"W).
Holotype male (LES 27734), Bahia, Feira da Mata, Gruna Boca da Lapa cave, 15 October 2020, leg. ME Bichuette, DF Torres, JS Gallo, L Senna-Horta & JE Gallão. Paratypes 1 male (parts in slides), 2 females (LES 27735); 1 male, 3 females (one female with parts in slides) (LES 27736), same data as holotype.
Body length: ♂ 8.0 mm, ♀ 9.3 mm. Body unpigmented, slightly robust, lateral sides slightly convex (Figs
Male. Pereopods 1–6 (Fig.
The new species is named after the locality where the specimens were collected, Gruna Boca da Lapa, an important cave discovered and mapped by Grupo Bambuí de Pesquisas Espeleológicas (GBPE) in 1991 and only now is it studied in relation to its cave fauna.
Xangoniscus lapaensis sp. nov. resembles X. aganju in the shape of the male pleopod 2 endopod; however, it differs in the vertex of the cephalon that lacks a frontal groove, and with a more pronounced suprantennal line that surpasses the antennule insertion (not surpassing in X. aganju). Also, the male pereopod 6 carpus lacks any modification (strongly modified in X. aganju, see
Xangoniscus lapaensis sp. nov. occurs in travertine pools fed by infiltration of water through the limestone rock, dripping from the cave ceiling, and in isolated pools along the cave (Fig.
Bahia, Feira da Mata, Gruna da Pingueira II cave (14°12'04.3"S, 44°24'39.1"W).
Holotype male (LES 27737), Bahia, Feira da Mata, Gruna da Pingueira II cave, 14 October 2020, leg. ME Bichuette, DF Torres, JS Gallo & JE Gallão. Paratypes 2 males, 3 females (one female with parts in micropreparations) (LES 27738), 6 males (one male in micropreparations), 4 females (LES 27739), same data as holotype.
Body length: ♂ 12 mm, ♀ 10 mm. Body unpigmented, slender and elongated, lateral sides almost parallel (Figs
Male. Pereopods 1–3 (Fig.
The new species is named after Alexandre Lobo for all his contributions for knowledge and conservation of the Brazilian speleological heritage.
Xangoniscus loboi sp. nov. differs from all other species of the genus in having the proximal lobe of male pereopods 1–3 merus covered with scales and fine setae, and in the shape of the distal complex apparatus of the male pleopod 2 endopod.
Xangoniscus loboi sp. nov. occurs in a small stream formed by infiltration water in the limestone rock (Fig.
Both species described herein show the classical signs of troglomorphism (eyes absent and body depigmented), as observed for the other eight valid species of Xangoniscus, i.e., X. aganju, X. ceci, X. dagua, X. ibiracatuensis, X. itacarambiensis, X. lundi, X. odara and X. santinhoi. This could be a pattern for the entire genus, since all species described until now show these same characteristics. Morphological variability in these two characters was not observed for any species of Xangoniscus. To date, no Xangoniscus species has been recorded outside caves, in the epigean environment, which may denote an ancient isolation of this group.
Morphologically, X. lapaensis sp. nov. and X. aganju (including specimens from Gruna Chico Pernambuco and Domingão caves, see
The Serra do Ramalho region has been locally exploited historically by agriculture activities and charcoal production on a small scale. At a broader scale, mining projects and large agriculture production along the boundaries of the region pose future threats (Gallão & Bichuette, 2018) (Fig.
A preliminary evaluation of the conservation status of these species was conducted following the IUCN (International Union of Conservation of Nature) classification. Despite their high population density and abundance, Xangoniscus lapaensis sp. nov. and Xangoniscus loboi sp. nov. were classified as Critically Endangered (CR) according to the criteria B2ab(iii). Both species are endemic, restricted to their type-locality and found in a specific microhabitat (travertine pools and streams formed by infiltration water). The surrounding areas of the cavities are impacted by deforestation for pastures and the native vegetation is present only at their entrances (Fig.
We thank ICMBio/CECAV (Instituto Chico Mendes para Conservação da Biodiversidade/Centro Nacional de Pesquisa e Conservação de Cavernas) for financial support to the project “Estudos para definição de áreas prioritárias para a Conservação de Proteção Espeleológico na Serra do Ramalho – BA”, under execution by the GBPE (Grupo Bambuí de Pesquisas Espeleológicas), coordinated by Ezio Rubbioli and integrated by MEB and LH; to the members of GBPE, E Rubbioli, F Chaimowicz, A Lobo and R Brandi for all support in the occasion of the discovery of the new species; a special thanks to GBPE for sharing speleological information of Serra do Ramalho karst area with MEB and for all contribution to the knowledge of Brazilian caves; to SISBIO (Sistema de Autorização Informação em Biodiversidade) for collection permit to MEB (28992); to Vicerrectoría de Investigaciones of the University of Cartagena for the financial support to the Grupo de Investigación en Biología Descriptiva y Aplicada (Project Redes de Investigación Científica, Acta de Compromiso N° 017–2020); to CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) for financial support to ISC-F during his postdoctoral fellowship (CAPES/PNPD/UFCG/CTRN/PPGRN/201713705) (Finance Code 001); to CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) for productivity fellowship to MEB (310378/2017–6). We would like to thank the anonymous reviewers and editor Dr. ML Niemiller for their critiques and suggestions for the work.