Corresponding author: Jairo A. Moreno-González (
Academic editor: J. A. Ochoa
We present a molecular phylogenetic analysis including a survey for overlooked phenotypic characters. Based on both analysis and characters a new cave-dwelling species is described:
Moreno-González JA, Pinto-da-Rocha R, Gallão JE (2021) Bringing order to a complex system: phenotypic and genotypic evidence contribute to the taxonomy of
Among the Neotropical buthid genera,
Great challenges need to be overcome before the taxonomy of
Species-level distinctions are also problematic in
The problems mentioned above, added to the fact that some species were described based on juveniles (e.g.,
There have been a few modern taxonomic revisions of
How to classify subterranean organisms based on their restriction/adaptation to the cave habitat has been a matter of debate for a long time (see
Arachnids (except
Troglobitic scorpions are globally rare (
In this contribution, we present a phylogenetic hypothesis including a survey for overlooked phenotypical characters. Based on both analysis and characters a new cave-dwelling species is described:
The type-material of the new species is housed in the Laboratório de Estudos Subterrâneos (
According to
Specimens were studied under a Leica MZ75 stereomicroscope with an ocular micrometer. Z-stack pictures under white light and UV light were taken using a Leica MC 170 HD camera. Habitus pictures were taken under white light using a Nikon D3300 digital camera and a 65 mm lens. For Scanning Electron Microscopy (SEM) imaging, a pectine was dissected and cleaned in distilled water with neutral detergent by ultrasound for one minute. After cleaning, the pectine was washed with distilled water and dehydrated via an ethanol concentration gradient (70%, 80%, 90%, 96%, and 100%), giving it 5–15 min in each concentration. Dehydration was completed under critical point drying with the pectine mounted onto a SEM stub using copper tape, after which it was sputter-coated with gold. Stubs were photographed using a Zeiss
General terminology follows
Pedipalp carinae:
Mesosoma, metasoma, and telson carinae:
Others:
The ingroup taxa comprised 31 terminals of 20 described species of
We extracted genomic DNA from leg tissues using the protocol of
List of primers used to amplify DNA sequences of
Locus | Primer | Sequences | Direction | Annealing (T, °C) | Reference |
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LCO1490-jj2 | 5’- CHA CWA AYC AYA ARG AYA TYG G | F | 49.3–62.0 |
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HCO2198-jj2 | 5’- ANA CTT CNG GRT GNC CAA ARA ATC A | R | 57.9–66.7 |
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12S | 12Sai | 5’- AAA CTA GGA TTA GAT ACC CTA TTA T | F | 52.3 |
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12S | 12Sbi | 5’- AAG AGC GAC GGG CGA TGT GT | R | 64.6 |
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12S | 12Sop2r | 5’ CCC TTA AAY YTA CTT TGT TAC GAC C | R | 50 |
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16S | 16Sbr | 5’- CTC CGG TTT GAA CTC AGA TCA | F | 57.7 |
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16S | 16S_F | 5’- CGA TTT GAA CTC AGA TCA | F | 49.3 |
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16S | 16Sbr_mod | 5’- GTG CAA AGG TAG CAT AAT CA | R | 53.7 |
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28S | 28Sa (Sad3) | 5’- GAC CCG TCT TGA AAC ACG GA | F | 60.3 |
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28S | 28Srd5b | 5’- CCA CAG CGC CAG TTC TGC TTA C | R | 64.2 |
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28S | 28SBout | 5’- CCC ACA GCG CCA GTT CTG CTT ACC | R | 68 |
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PCR amplifications were checked using electrophoresis of agarose gel (2% agarose). Positive amplifications were purified using Agencourt Ampure XP (Beckman Coulter), then quantified using a Thermo Scientific NanoDrop spectrophotometer. We prepared sequencing reactions with the BigDye Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems), precipitated PCR products with sodium acetate, and sequenced using an ABI PRISM 3100 Genetic Analyzer/HITACHI (Applied Biosystems). Sequence editing (e.g., primer trimming) and contiguous sequence generation were made on AB1 files using Geneious R11 (
Ribosomal gene and intron sequences (12S rDNA, 16S rDNA, and 28S rDNA) were aligned using the E-INS-i algorithm of MAFFT (
List of terminals, voucher specimens, and sequences (GenBank accession numbers indicated) used in the phylogenetic analysis of
Species | Subgenus | Voucher | 12S | 16S | 28S |
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MACN Ar 35705 |
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MACN Ar 35713 |
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MACN Ar 35708 |
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MACN Ar 35709 |
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MACN Ar 35693 |
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MACN Ar 35694 |
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MACN Ar 35695 |
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MACN Ar 35723 |
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MACN Ar 35724 |
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MACN Ar 35712 |
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MACN Ar 35714 |
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MACN Ar 35715 |
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MACN Ar 35716 |
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MACN Ar 35717 |
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MACN Ar 35718 |
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Tree search was conducted in IQTREE using the maximum likelihood (
The tree log-likelihood score was -12896.086. The best-fit models per molecular partition were TIM2+F+G4 (12S), TIM2+F+I+G4 (16S), TNe+R2 (28S), and TIM+F+I+G4 (
Phylogeny of
Phylogeny of
Phylogeny of
We observed that the distribution of the ventral setae of telotarsi I–IV in
Phenotypic characters useful for the taxonomy of
Species | Subgenus | Species Group | Telotarsal setae | Females | |
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Type II | Absent |
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Type I | Absent | D= semicircular | |
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Type II | First 2/3 of the anterior region |
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Type II | More than first 2/3 of the anterior region |
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Type II | First 2/3 of the anterior region |
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Type II | First 2/3 of the anterior region |
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Type II | First 2/3 of the anterior region |
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Type II | First 2/3 of the anterior region |
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Type I | First anteromedian third |
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Type II | First 2/3 of the anterior region |
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Type II | Absent | D= suboval | |
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Type II | Absent | D= suboval | |
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Type I | Absent | D= semicircular | |
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Type II | Absent | D= semicircular | |
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Type II | First 2/3 of the anterior region |
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Type II | Absent | D= semicircular | |
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Type II | First anteromedian third |
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? | Absent | D= semicircular | |
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Type II | More than first 2/3 of the anterior region |
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Type II | First anteromedian third |
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Type II | First 2/3 of the anterior region |
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Type I | Absent | D= semicircular |
The pectinal basal piece of female exhibits the following character states within the examined terminals of
However, it is noteworthy that in some
This subgenus currently includes, among others, all species assigned to the
Brazil: State of Goiás:
The species epithet is a derivative form of the Greek noun, σπήλαιον (Latin: caverna), which means cave, in reference to the subterranean habitat where
(Based on female). This species belongs to the
On the other hand,
Finally,
In an unpublished comprehensive phylogenetic analysis of
On the other hand,
Based on the female holotype (
Total length. Female: 53.52 mm (measurements in Table
Measurements (mm) of
Structure | Measure | Female holotype | Female paratype # 1 | Female paratype # 2 | Female paratype #4 | Female paratype #5 | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype | Female paratype |
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LES014668 | LES014669 | LES014670 | LES014671 | LES014672 | LES014673 | LES014673 | LES014673 | |||||||||||
Total length | – | 53.52 | 51.29 | 57.89 | 51.06 | 51.70 | 48.30 | 49.64 | 51.41 | 49.90 | 57.98 | 53.69 | 53.84 | 54.19 | 54.40 | 50.75 | 52.02 | 54.29 |
Carapace | length | 6.00 | 5.84 | 6.40 | 5.68 | 5.68 | 5.57 | 5.57 | 5.71 | 5.57 | 6.45 | 5.83 | 5.98 | 6.06 | 6.20 | 5.57 | 5.73 | 6.07 |
Carapace | anterior width | 4.08 | 3.76 | 4.32 | 3.84 | 3.84 | 3.71 | 3.57 | 3.86 | 3.71 | 3.33 | 3.08 | 3.09 | 3.13 | 3.28 | 2.89 | 3.01 | 3.10 |
Carapace | posterior width | 6.64 | 6.48 | 7.12 | 6.24 | 6.24 | 6.00 | 5.57 | 6.29 | 5.71 | 6.53 | 6.09 | 6.08 | 6.10 | 6.32 | 5.84 | 5.92 | 6.21 |
Carapace | eye diameter | 0.45 | 0.48 | 0.48 | 0.45 | 0.45 | 0.40 | 0.40 | 0.43 | 0.47 | 0.47 | 0.44 | 0.44 | 0.46 | 0.43 | 0.40 | 0.41 | 0.43 |
Carapace | interocular distance | 0.53 | 0.50 | 0.55 | 0.48 | 0.48 | 0.47 | 0.47 | 0.50 | 0.53 | 0.54 | 0.53 | 0.49 | 0.54 | 0.55 | 0.54 | 0.47 | 0.59 |
Carapace | ocular diada width | 1.20 | 1.28 | 1.36 | 1.20 | 1.20 | 1.13 | 1.17 | 1.20 | 1.17 | 1.26 | 1.21 | 1.24 | 1.21 | 1.23 | 1.13 | 1.18 | 1.19 |
Tergite I | length | 1.12 | 1.08 | 1.08 | 1.00 | 1.00 | 1.00 | 1.07 | 1.00 | 0.87 | 1.24 | 1.16 | 1.18 | 1.16 | 1.14 | 1.09 | 1.01 | 1.17 |
Tergite II | length | 1.44 | 1.28 | 1.52 | 1.24 | 1.24 | 1.27 | 1.27 | 1.27 | 1.00 | 1.50 | 1.32 | 1.44 | 1.42 | 1.35 | 1.23 | 1.31 | 1.41 |
Tergite III | length | 1.76 | 1.60 | 1.88 | 1.56 | 1.68 | 1.47 | 1.73 | 1.60 | 1.67 | 1.88 | 1.77 | 1.68 | 1.61 | 1.68 | 1.64 | 1.67 | 1.76 |
Tergite IV | length | 2.20 | 2.00 | 2.32 | 2.08 | 2.04 | 1.87 | 2.00 | 2.07 | 1.93 | 2.35 | 2.16 | 2.20 | 2.01 | 2.10 | 1.94 | 2.00 | 2.06 |
Tergite V | length | 2.32 | 2.20 | 2.60 | 2.28 | 2.24 | 2.07 | 2.20 | 2.20 | 2.33 | 2.52 | 2.44 | 2.34 | 2.33 | 2.28 | 2.19 | 2.28 | 2.38 |
Tergite |
length | 2.68 | 2.40 | 2.84 | 2.44 | 2.52 | 2.27 | 2.40 | 2.40 | 2.53 | 2.74 | 2.56 | 2.58 | 2.48 | 2.51 | 2.44 | 2.47 | 2.56 |
Tergite VII | length | 3.88 | 3.96 | 4.20 | 3.80 | 3.84 | 3.60 | 3.80 | 4.07 | 3.80 | 4.60 | 4.09 | 4.06 | 4.05 | 4.11 | 4.01 | 3.83 | 4.28 |
Mesosoma | total length (tergites) | 15.40 | 14.52 | 16.44 | 14.40 | 14.56 | 13.53 | 14.47 | 14.60 | 14.13 | 16.83 | 15.50 | 15.48 | 15.06 | 15.17 | 14.54 | 14.57 | 15.62 |
Metasoma I | length | 3.55 | 3.55 | 3.75 | 3.55 | 3.55 | 3.30 | 3.50 | 3.60 | 3.50 | 4.00 | 3.67 | 3.63 | 3.66 | 3.71 | 3.49 | 3.56 | 3.84 |
Metasoma I | width | 2.85 | 2.85 | 2.95 | 2.35 | 2.85 | 2.50 | 2.70 | 2.90 | 2.70 | 2.99 | 2.83 | 2.79 | 2.88 | 2.86 | 2.78 | 2.76 | 2.85 |
Metasoma I | height | 2.60 | 2.65 | 2.65 | 2.85 | 2.85 | 2.40 | 2.50 | 2.50 | 2.50 | 2.83 | 2.62 | 2.62 | 2.66 | 2.70 | 2.43 | 2.50 | 2.63 |
Metasoma II | length | 4.55 | 4.40 | 4.80 | 4.35 | 4.50 | 4.20 | 4.30 | 4.50 | 4.30 | 4.97 | 4.57 | 4.67 | 4.69 | 4.71 | 4.32 | 4.57 | 4.64 |
Metasoma II | width | 2.75 | 2.70 | 3.05 | 2.20 | 2.75 | 2.40 | 2.60 | 2.70 | 2.60 | 3.02 | 2.75 | 2.82 | 2.84 | 2.89 | 2.69 | 2.72 | 2.77 |
Metasoma II | height | 2.75 | 2.70 | 2.95 | 2.70 | 2.85 | 2.40 | 2.50 | 2.60 | 2.50 | 2.73 | 2.54 | 2.45 | 2.51 | 2.59 | 2.44 | 2.48 | 2.60 |
Metasoma III | length | 5.20 | 5.00 | 5.50 | 4.90 | 5.00 | 4.70 | 4.80 | 5.00 | 4.90 | 5.44 | 5.19 | 5.10 | 5.28 | 5.21 | 4.68 | 4.96 | 5.05 |
Metasoma III | width | 2.85 | 2.70 | 3.20 | 2.25 | 2.80 | 2.40 | 2.70 | 2.70 | 2.60 | 3.09 | 2.77 | 2.79 | 2.83 | 2.88 | 2.67 | 2.79 | 2.95 |
Metasoma III | height | 2.90 | 2.60 | 3.00 | 2.85 | 2.85 | 2.40 | 2.40 | 2.50 | 2.60 | 2.81 | 2.53 | 2.54 | 2.60 | 2.63 | 2.52 | 2.50 | 2.58 |
Metasoma IV | length | 5.75 | 5.50 | 6.50 | 5.50 | 5.60 | 5.20 | 5.40 | 5.60 | 5.50 | 6.25 | 5.88 | 5.81 | 5.80 | 5.87 | 5.51 | 5.79 | 5.82 |
Metasoma IV | width | 2.85 | 2.75 | 3.25 | 2.25 | 2.85 | 2.50 | 2.60 | 2.80 | 2.60 | 3.10 | 2.77 | 2.77 | 2.87 | 2.87 | 2.68 | 2.77 | 2.86 |
Metasoma IV | height | 2.85 | 2.50 | 3.12 | 2.85 | 2.85 | 2.30 | 2.40 | 2.50 | 2.40 | 2.77 | 2.55 | 2.51 | 2.49 | 2.64 | 2.59 | 2.51 | 2.58 |
Metasoma V | length | 6.83 | 6.57 | 7.74 | 6.70 | 6.76 | 6.00 | 6.10 | 6.50 | 6.30 | 7.24 | 6.74 | 6.67 | 6.93 | 6.92 | 6.43 | 6.58 | 6.84 |
Metasoma V | width | 2.93 | 2.60 | 3.25 | 2.86 | 2.93 | 2.30 | 2.50 | 2.60 | 2.40 | 2.91 | 2.57 | 2.58 | 2.59 | 2.63 | 2.49 | 2.57 | 2.59 |
Metasoma V | height | 2.73 | 2.54 | 3.12 | 2.80 | 2.86 | 2.30 | 2.40 | 2.60 | 2.40 | 2.69 | 2.82 | 2.50 | 2.48 | 2.58 | 2.41 | 2.51 | 2.61 |
Metasoma | length | 25.88 | 25.02 | 28.29 | 25.00 | 25.41 | 18.70 | 19.30 | 20.20 | 19.60 | 27.90 | 26.05 | 25.88 | 26.36 | 26.42 | 24.43 | 25.46 | 26.19 |
Telson | vesicle length | 3.84 | 3.77 | 4.23 | 3.77 | 3.77 | 3.40 | 3.40 | 3.70 | 3.50 | 3.57 | 3.41 | 3.38 | 3.46 | 3.51 | 3.33 | 3.37 | 3.43 |
Telson | vesicle width | 2.21 | 1.95 | 2.28 | 1.95 | 1.95 | 1.80 | 1.90 | 2.00 | 1.90 | 2.10 | 1.96 | 1.93 | 2.05 | 2.11 | 1.88 | 1.89 | 2.03 |
Telson | vesicle height | 2.15 | 2.08 | 2.28 | 2.08 | 2.02 | 1.80 | 1.90 | 2.00 | 1.90 | 2.17 | 2.01 | 2.00 | 2.10 | 2.12 | 1.90 | 1.97 | 2.11 |
Telson | aculeus length | 2.80 | 2.67 | 2.99 | 2.67 | 2.67 | 2.50 | 2.60 | 2.70 | 2.60 | 2.68 | 2.54 | 2.53 | 2.63 | 2.56 | 2.52 | 2.49 | 2.57 |
Telson | total length | 6.24 | 5.92 | 6.76 | 5.98 | 6.05 | 5.80 | 5.50 | 5.90 | 5.70 | 6.80 | 6.31 | 6.50 | 6.71 | 6.61 | 6.21 | 6.26 | 6.41 |
Metasoma+ Telson | total length | 32.12 | 30.93 | 35.05 | 30.98 | 31.46 | 29.20 | 29.60 | 31.10 | 30.20 | 34.70 | 32.36 | 32.38 | 33.07 | 33.03 | 30.64 | 31.72 | 32.60 |
Femur | length | 6.18 | 5.98 | 6.70 | 6.18 | 6.11 | 5.60 | 5.80 | 6.00 | 5.90 | 6.96 | 6.46 | 6.51 | 6.76 | 6.72 | 6.39 | 6.41 | 6.61 |
Femur | width | 1.50 | 1.56 | 1.76 | 1.69 | 1.56 | 1.40 | 1.50 | 1.50 | 1.50 | 1.75 | 1.56 | 1.61 | 1.62 | 1.64 | 1.54 | 1.60 | 1.60 |
Patella | length | 6.76 | 6.70 | 7.28 | 6.44 | 6.57 | 6.00 | 6.00 | 6.20 | 6.30 | 6.68 | 6.10 | 6.12 | 6.54 | 6.31 | 5.78 | 6.01 | 6.21 |
Patella | width | 2.08 | 2.02 | 2.02 | 2.08 | 2.02 | 1.80 | 1.90 | 2.00 | 1.90 | 2.12 | 1.88 | 1.93 | 2.01 | 2.00 | 1.85 | 1.90 | 1.94 |
Chela | length | 11.50 | 11.10 | 13.00 | 11.57 | 10.50 | 10.40 | 10.60 | 11.20 | 10.90 | 12.26 | 11.28 | 11.25 | 11.75 | 11.77 | 10.79 | 11.08 | 11.63 |
Chela | width | 2.60 | 2.00 | 2.47 | 2.34 | 2.10 | 1.80 | 2.00 | 2.00 | 1.90 | 2.32 | 2.04 | 1.96 | 2.06 | 2.08 | 1.90 | 2.01 | 2.02 |
Chela | height | 2.10 | 2.10 | 2.73 | 2.21 | 2.10 | 1.80 | 2.00 | 2.10 | 2.00 | 2.10 | 1.81 | 1.85 | 1.92 | 1.95 | 1.84 | 1.82 | 1.93 |
Chela | movable finger length | 8.00 | 7.20 | 8.97 | 7.80 | 7.20 | 7.00 | 7.00 | 7.40 | 7.30 | 8.29 | 7.62 | 7.61 | 8.01 | 8.02 | 7.45 | 7.51 | 7.99 |
Chela | fixed finger length | 6.80 | 6.40 | 7.67 | 6.89 | 6.20 | 6.20 | 6.10 | 6.60 | 6.40 | 7.07 | 6.55 | 6.74 | 7.05 | 6.91 | 6.17 | 6.89 | 6.92 |
Chela | palm length | 4.00 | 3.80 | 4.68 | 4.16 | 3.90 | 3.40 | 3.70 | 3.70 | 3.70 | 4.14 | 4.11 | 4.09 | 4.16 | 4.15 | 3.92 | 4.03 | 4.03 |
General pattern (in ethanol 70%) (Fig.
Variation in the number of macrosetae of the ventrosubmedian setal rows on telotarsi I–IV across paratypes of
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8/7 | 8/7 | 6/7 | 8/6 | 8/7 | 7/8 | 6/7 | 6/7 |
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7/8 | 8/7 | 8/8 | 8/9 | 8/8 | 7/8 | 8/8 | 8/7 |
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8/8 | 7/8 | 8/7 | 8/9 | 10/8 | 8/9 | 6/8 | 8/7 |
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10/10 | 10/10 | 10/10 | - | 9/12 | 10/9 | 10/10 | 10/11 |
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6/6 | - | 7/8 | 7/6 | 7/6 | 7/7 | 6/7 | 6/8 |
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7/7 | 7/8 | 7/7 | 8/7 | 9/6 | 7/7 | 8/7 | 7/8 |
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9/7 | 8/8 | 7/8 | 8/7 | 7/6 | 10/8 | 6/6 | 8/8 |
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- | 10/12 | 10/11 | 9/8 | 10/10 | 11/10 | 10/8 | 10/10 |
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8/7 | 8/7 | 8/9 | 8/8 | 9/8 | 6/7 | 8/8 | 9/9 |
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8/7 | 8/7 | 10/9 | 9/8 | 9/8 | 10/7 | 9/9 | 8/8 |
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8/8 | 9/8 | 9/10 | 9/9 | 9/8 | 8/8 | 9/9 | 9/9 |
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11/11 | 11/10 | 10/11 | 10/11 | 10/10 | 10/10 | 12/12 | 10/11 |
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9/8 | 8/8 | 8/8 | 8/8 | 8/9 | 9/8 | 8/8 | 9/9 |
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9/8 | 8/8 | 8/9 | 8/8 | 10/9 | 9/8 | 8/8 | 8/8 |
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9/9 | 10/9 | 9/9 | 8/8 | 9/9 | 10/10 | 9/9 | 9/8 |
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10/11 | 11/12 | 11/10 | 11/10 | 12/12 | 11/12 | 12/11 | 11/12 |
Russão II cave is formed by limestone (a karstified type of rock), located in Posse municipality, the northeastern state of Goiás, Central Brazil. This karst region is part of the Bambuí geomorphological group, the large geomorphological group in Brazil, occurring in states of Bahia, Goiás, Minas Gerais, and Tocantins. Russão II cave is inserted on the Cerrado morphoclimatic domain (
This species was studied in the past by
Habitat of
Our phylogenetic results (Figs
More recently,
Finally, other aspects were not challenged by several authors, such as the consensus about
In the original description of
It is worth mentioning that the slender and elongated shape of the male pedipalp has been demonstrated to be a highly homoplastic character state that evolved independently at least four times within
For example, the position of
Given the results of our phylogenetic analysis, previous hypotheses (
The leg telotarsi ventral setation has been a very commonly used phenotypic character to define genera and/or assist species diagnoses in families such as
It is particularly interesting to note that the
This previously ignored morphological character has sometimes been proved useful to assist taxonomic delimitations, even outside the genus
The sexual dimorphism of the basal pectinal piece and the glands that it sometimes carries are characters that have been neglected in the taxonomy of
The glandular region of the pectinal basal piece of female has far too often been an overlooked morphological character in taxonomic and systematic contributions dealing with buthid taxa.
On the other hand, the dilatation exhibited by the basal middle lamellae of the female pectines has been a widely used character in the taxonomy of
Cuticular (exocrine) glandular regions are a very common feature in a broad spectrum of arthropod groups (e.g.,
Species of two scorpion families occur in Brazilian caves,
Representatives of
As expected, troglophilic populations are found more often inside caves than in epigean habitats due to differences in the dynamics of species. They are generally more numerous in subterranean habitats (
Troglophiles and trogloxenes are both found in epigean and subterranean environments, and, since individuals can move between them, it is not easy to distinguish between these two categories. One strong piece of evidence for troglophilic populations is the presence of individuals of all ages distributed along with the subterranean environment throughout different annual cycles (
No individuals of
Also, it is worth mentioning that troglophiles are not less adapted to subterranean environment than troglobites in what is considered a continuum of cave adaptation (
We are grateful to Alex S. Valdarnini (Grupo Pierre Martin de Espeleologia – GPME) for the photography of Russão II cave. Eleonora Trajano and Maria E. Bichuette (
Voucher samples from which material was examined for morphological study and Sanger loci were sequenced. The vouchers are deposited in the following collections: the Ambrose Monell Cryocollection (