TYPE Original Research

PUBLISHED 15 June 2023

DOI 10.3389/fevo.2023.1177627

Phylogenomics of
OPEN ACCESS Lasiodoriforms: reclassification of
the South American genus Vitalius
EDITED BY
Felipe Gobbi Grazziotin,
Instituto Butantan, Brazil

REVIEWED BY
Marc Domènech,
Lucas, Silva and Bertani and allied
University of Bristol, United Kingdom
Jesus Lozano-Fernandez,
University of Barcelona, Spain, in collaboration
genera (Araneae: Theraphosidae)
with reviewer MD
Carlos Perafán,
University of Caldas, Colombia Arthur Galleti-Lima1*, Chris A. Hamilton2 , Leandro M. Borges3 and
*CORRESPONDENCE José Paulo L. Guadanucci1
Arthur Galleti-Lima
1
galleti.lima@unesp.br Departamento de Biodiversidade, Instituto de Biociências, Universidade Estadual Paulista “Júlio de
Mesquita Filho” (UNESP), São Paulo, Brazil, 2 Department of Entomology, Plant Pathology and
RECEIVED 01 March 2023 Nematology, University of Idaho, Moscow, ID, United States, 3 Pós-Graduação em Biodiversidade Animal,
ACCEPTED 24 May 2023 Universidade Federal de Santa Maria, Santa Maria, Rio Grande do Sul, Brazil
PUBLISHED 15 June 2023

CITATION
Galleti-Lima A, Hamilton CA, Borges LM and
Theraphosinae is the most diverse subfamily of Theraphosidae spiders, but their
Guadanucci JPL (2023) Phylogenomics of
Lasiodoriforms: reclassification of the South evolutionary history remains unresolved to date. This problem is common in
American genus Vitalius Lucas, Silva and Bertani taxonomic groups with phylogenetic hypotheses that have often been based
and allied genera (Araneae: Theraphosidae).
only on qualitative morphological characters and, rarely, on molecular analyses.
Front. Ecol. Evol. 11:1177627.
doi: 10.3389/fevo.2023.1177627 Phylogenomics has significantly contributed to the understanding of the evolution
COPYRIGHT
of many non-model groups, such as spiders. Herein, we employed ultraconserved
© 2023 Galleti-Lima, Hamilton, Borges and elements (UCEs) phylogenomics to propose a new hypothesis for a group of
Guadanucci. This is an open-access article Theraphosinae genera, namely Lasiodoriforms: Vitalius, Lasiodora, Eupalaestrus,
distributed under the terms of the Creative
Commons Attribution License (CC BY). The use,
Pterinopelma, Proshapalopus, and Nhandu. We propose three genera and their
distribution or reproduction in other forums is respective morphological diagnoses are provided. Our phylogeny supports the
permitted, provided the original author(s) and transfer of species from the genus Vitalius to Pterinopelma and Proshapalopus
the copyright owner(s) are credited and that
the original publication in this journal is cited, in
and from Proshapalopus to Eupalaestrus. Finally, we describe a new species of
accordance with accepted academic practice. Vitalius from Southern Brazil. Based on these three new generic descriptions
No use, distribution or reproduction is and transferred species, the Lasiodoriform tarantulas comprise nine genera from
permitted which does not comply with these
terms.
Argentina, Brazil, Paraguay, and Uruguay, and the genus Vitalius now includes
seven species.

KEYWORDS

Mygalomorphae, Theraphosidae, new world species, phylogenomics, Taxonomy

1. Introduction
The subfamily Theraphosinae is endemic to the Neotropics and is one of twelve
currently included in Theraphosidae Thorell, 1896, the family that includes tarantulas
(Guadanucci, 2014; Lüddecke et al., 2018; Turner et al., 2018; Foley et al., 2019).
Theraphosinae comprises the greatest diversity within the family, currently including 69
genera and more than 500 species (World Spider Catalog, 2023), distributed mainly in
the Neotropics (Central America, South America, southern Mexico, and the Caribbean),
as well as in the southwestern United States and northern Mexico temperate regions
(Hamilton et al., 2016b; Mendoza and Francke, 2017; Pérez-Miles and Perafán, 2020).

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Galleti-Lima et al.
Defining species boundaries and identifying useful
morphological characters are challenging tasks when dealing
with theraphosids, as they present homogeneous morphology
and continuous variation across genera and species (Bertani,
2001; Fukushima and Bertani, 2017). Phylogenetic reconstructions
from morphological matrices often result in low-supported clades
and trees with many homoplastic characters and, consequently,
discordant evolutionary hypotheses. In addition, morphological
similarities are even greater in females, making it difficult to
identify genera and species (Bertani et al., 2011, 2012; Bertani and
Leal, 2016; Hamilton et al., 2016b; Galleti-Lima and Guadanucci,
2018; Perafán and Valencia-Cuellar, 2018).
When taking a broader look at informative morphological
characters, there have been a few hypotheses among Theraphosidae
subfamilies that have shown that the monophyly of Theraphosinae
is supported by the following synapomorphies: keels on the
male palpal bulb, the subtegulum extended over the tegulum,
and the presence of different types of urticating setae (types
I, III, IV, V, VI, or VII) (Raven, 1985; Pérez-Miles et al., 1996;
Bertani and Guadanucci, 2013; Guadanucci, 2014; Kaderka
et al., 2019). This has also been shown as monophyletic
in molecular analyses (Lüddecke et al., 2018; Foley et al.,
2019). Additionally, there have been hypotheses based on
the morphological features that proposed to explain the
interrelationships of theraphosinae genera (Pérez-Miles et al.,
1996; Bertani, 2001; Bertani et al., 2011; Bertani and Guadanucci,
2013; Perafán et al., 2015; Galleti-Lima and Guadanucci, 2018;
Perafán and Valencia-Cuellar, 2018; Fabiano-Da-Silva et al.,
2019), as well as one phylogenetic hypothesis using molecular
data, based on a single mitochondrial gene region (Turner et al.,
2018).
Despite disagreements between studies, some phylogenetic
lineages within Theraphosinae have been recovered as
monophyletic in both morphological and molecular hypotheses.
For example, clades supported by the presence of different types
of urticating setae (III + IV and III + I) (Pérez-Miles et al., 1996;
Turner et al., 2018), the “spoon-like” embolus clade (Bertani,
2001), and a group supported by the presence of subapical and
retrolateral keels in the male palpal bulb (Bertani, 2001; Turner
et al., 2018). In their phylogenetic hypothesis, Turner et al.
(2018) recommended the division of Theraphosinae into three
tribes: Grammostolini, tarantulas with type IV urticating setae;
Hapalopini, lineage where most genera possess only type III
urticating setae also known as “dwarf tarantulas” and the sister
lineage of Grammostolini; and Theraphosini, tarantulas with type
I urticating setae, such as Vitalius Lucas et al., 1993, and its closely
related genera.
The group of tarantulas that share the presence of subapical
and retrolateral keels in the male palpal bulb include the following
South American genera: Lasiodora Koch, 1850, Eupalaestrus
Pocock, 1901, Pterinopelma Pocock, 1901, Proshapalopus Mello-
Leitão, 1923, Nhandu Lucas, 1983, and Vitalius. It has been
recovered as monophyletic in recent phylogenies (Bertani,
2001; Bertani et al., 2012; Galleti-Lima and Guadanucci, 2018;
Perafán and Valencia-Cuellar, 2018; Turner et al., 2018). The
first study to focus on these tarantulas, carried out by Bertani
(2001), produced a phylogenetic hypothesis using morphological
Frontiers in Ecology and Evolution
10.3389/fevo.2023.1177627
characteristics and a taxonomic revision of Eupalaestrus,
Prohapalopus, Nhandu, and Vitalius. Intrageneric relationships
within Vitalius have never been fully resolved, with most species
forming a poorly solved polytomic clade (Bertani, 2001; Bertani
et al., 2011; Galleti-Lima and Guadanucci, 2018; Perafán and
Valencia-Cuellar, 2018), and in some topologies, the genus was
not recovered as monophyletic (Perafán and Valencia-Cuellar,
2018).
More recently, Bertani et al. (2011) proposed the revalidation
of Pterinopelma, with P. vitiosum (Keyserling, 1891) as the type-
species, and provided a description of P. sazimai (Bertani et al.,
2011). In the subsequent year, Bertani et al. (2012) proposed the
transfer of Vitalius nondescriptus (Mello-Leitão, 1926), which is
sympatric with P. sazimai, and in 2016, Bertani and Leal (2016)
described Pterinopelma felipeleitei and the male of P. sazimai,
which was considered as a misidentification of the male of P.
felipeleitei by Bertani et al. (2011). In addition, the authors discussed
that P. sazimai perhaps should not be included in Pterinopelma
due to the morphology of the male palpal bulb. The monophyly
of the genus Pterinopelma, its interrelationships, and species
composition are key points for understanding the phylogeny of
Vitalius and Nhandu. In a phylogenetic study that aimed to
understand the evolution of stridulating setae in Theraphosinae
spiders (Galleti-Lima and Guadanucci, 2018), Pterinopelma was
not recovered as monophyletic, and the species P. sazimai was
closely related to Nhandu and distantly related from other
Pterinopelma species.
New molecular sequencing techniques can be used to access
hundreds or thousands of loci for phylogenetic inference. Targeted
enrichment approaches (e.g., Ultraconserved Elements and
Anchored Hybrid Enrichment) (Faircloth et al., 2012; Lemmon and
Lemmon, 2013) have been useful for inferring robust phylogenetic
relationships of the non-model groups, such as Mygalomorphae
spiders, such as Aphonopelma Pocock, 1901, Aptostichus
Simon, 1891, Aliatypus Smith, 1908, Antrodiaetus Ausserer,
1871, Hexurella Gertsch and Platnick, 1979, Mecicobothrium
Holmberg, 1882, Megahexura Kaston, 1972, Porrhothele Simon,
1892 (Faircloth et al., 2012; Hamilton et al., 2016b; Starrett
et al., 2016; Yeates et al., 2016; Branstetter et al., 2017; Hedin
et al., 2018, 2019; Derkarabetian et al., 2019; Zhang et al.,
2019; Kulkarni et al., 2020; Xu et al., 2021). To date, there
has been only one study using either of these phylogenomic
approaches in tarantulas, the North American genus Aphonopelma
(Hamilton et al., 2016a). Kulkarni et al. (2020) designed a
spider-specific UCE probe set, which has been recovering a high
number of loci, contributing to several questions in different
phylogenetic studies.
Motivated by the need for a robust phylogenetic
hypothesis that includes all diversity of Vitalius, Nhandu,
and Pterinopelma to support taxonomic reclassifications,
we present the first phylogenomic study of Theraphosinae
spiders in South America using sequences of UCEs. In
addition to the representatives of Lasiodora, Eupalaestrus,
Pterinopelma, Proshapalopus, Nhandu, and Vitalius, we
tested the monophyly of Vitalius, including all species, and
propose reclassifications for some of these genera based on the
resulting topology.
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2. Materials and methods
2.1. Molecular taxon sampling
All Theraphosinae species used in this research are deposited in
the Coleção Aracnológica Diamantina (CAD). A list with a voucher
number of these specimens is assembled in Supplementary File S1.
All type-species from genera belonging to the South American
lineage that possesses bulbs with retrolateral and subapical keels
were included in the analysis. Because we were seeking to
resolve the polytomies of the past morphological phylogenetic
hypotheses, all Vitalius species were included. For outgroup
comparison, we used morphologically distinct theraphosine genera
from ingroup taxa (i.e., without subapical and retrolateral
keels), such as Aphonopelma hentzi (Girard, 1853), Cyrtopholis
portoricae Chamberlin, 1917, an unidentified Sericopelma Ausserer,
1875, and Stichoplastoris Rudloff, 1997. All taxonomic decisions
were made based on node supports (>90) and diagnosable
morphological differences.
2.2. Morphology
Images and measurements used in morphological descriptions
were obtained with a Leica M205 C stereo microscope and the Leica
Application Suite V4.12 software. In the description of the new
species, body length measurements include the chelicerae, but not
the spinnerets. Leg segment lengths were measured in the dorsal
view between the joints. The carapace, eye tubercle, labium, and
sternum were measured in length and width. We followed Bertani
(2001) for the use of the terminologies of structures and spination.
The specimens used for images and descriptions are deposited
in the following collections: CAD, Coleção Aracnológica
Diamantina, Universidade Estadual Paulista, Rio Claro, Brazil—J.
P. L. Guadanucci; IBSP, Instituto Butantan, São Paulo, Brazil—A.
D. Brescovit; MCN, Museu de Ciências Naturais, Fundação
Zoobotânica do Rio Grande do Sul, Porto Alegre, Brazil—R. Ott;
MCTP, Museu de Ciências e Tecnologia, Pontifícia Universidade
Católica do Rio Grande do Sul, Porto Alegre, Brazil—R. A.
Teixeira; MZSP, Museu de Zoologia da Universidade de São Paulo,
São Paulo, Brazil—R. Pinto da Rocha; and UFMG, Universidade
Federal de Minas Gerais, Belo Horizonte, Brazil—A. Santos.
2.3. DNA extraction and phylogenomics
Genomic DNA was extracted from muscle tissues using the
Qiagen DNeasy Tissue KitTM (Qiagen, Valencia, CA, USA)
or OmniprepTM (G-Biosciences) and then quantitatively and
qualitatively assessed on agarose gel electrophoresis and a Qubit
2.0 Fluorometer (Thermo Fisher Scientific). Library prep, UCE
hybridization, and high-throughput Illumina sequencing were
carried out at Rapid Genomics (Gainesville, FL). DNA was sheared
to a mean fragment length of 500 bp, fragments were end-repaired
and A-tailed, followed by the incorporation of unique dual-indexed
Illumina adaptors and PCR enrichment. Samples were equimolar
pooled and sequenced on an SP flow cell (2 × 250 bp) or an
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10.3389/fevo.2023.1177627
S4 flow cell (2 × 150 bp). The resulting data were processed
using Phyluce version 1.7.1 (Faircloth, 2016) and a combined
arachnid-spider hybrid probeset (Starrett et al., 2016; Kulkarni
et al., 2020), where match settings for minimum identity and
minimum coverage of 85 and 85 (respectively) were used. Loci
were then aligned using MAFFT (Katoh and Standley, 2013),
with alignments externally trimmed and then internally trimmed
using gblocks with b1, b2, b3, and b4 and settings of 0.5, 0.5,
5, and 10, respectively, to remove poorly aligned blocks within
the sequences. The data were then additionally cleaned using
AMAS v 1.0 (Borowiec, 2016) and SPRUCEUP (Borowiec, 2019),
which removes poorly aligned sequence fragments from individual
sequences within alignments. Visual examination of distance
distribution plots identified a setting of 0.9 as optimal for trimming,
with a 50% occupancy matrix then being generated for use in
subsequent analyses. A maximum likelihood-based phylogeny was
inferred from 1,375 loci using IQTree2 (Nguyen et al., 2015;
Minh et al., 2020), with nodal support determined by the ultrafast
bootstrap and the SH-aLRT test (Anisimova et al., 2011) for
1,000 replicates each. All 1,375 loci were modeled independently
(for details on each model used, see “brazil.0.9.iqtree.log” file
attached in Supplementary File S2). All analyses were performed
on the University of Idaho Research Computing and Data Services
(RCDS) high-performance computing cluster. Tree edits were
performed using FigTree V. 1.4.4 software (Rambaut, 2018)
and then edited a image editor. DNA sequence alignments and
associated phylogenetic trees and data matrices, accompanying tree
files, and scripts have been deposited in Supplementary File S2.
The updated new names were used in the terminal taxa
of the Lasiodoriforms phylogenetic tree. Consider the prefix of
the specimens in the tree as referring to the genus to which
the specimen previously belonged (e.g., PROS, Proshapalopus;
EUPA, Eupalaestrus; LASI, Lasiodora; PTER, Pterinopelma; NHAN,
Nhandu; VITA, Vitalius).
3. Results
3.1. Phylogeny
The dataset comprised 1,375 loci (1,194,888 base pairs) with
63 terminals (6 outgroup and 57 ingroup terminals). The resulting
trees from the ML analysis are shown at the genera level
(Figure 1) and species level (Figure 2). The clade named herein as
Lasiodoriforms was recovered and well-supported (>90) by both
the SH-aLRT test and ultra-fast bootstrapping. The support values
in the nodes of each genus were also high in both tests, except
for the node referring to Proshapalopus multicuspidatus + Vitalius
nondescriptus. It is important to note that long branches of certain
lineages (e.g., P. multicuspidatus and P. felipeleitei) are because of
data quality issues (i.e., gaps and missing data). The accentuated
branch length is not likely to be a true representation—though the
phylogenetic placement of these lineages do not appear to change
following significant data cleaning. Our concatenated analysis
indicated Eupalaestrus (including P. amazonicus) as a sister group
of the remaining Lasiodoriforms. The node at the evolutionary
divergence between P. multicuspidatus + V. nondescriptus and the
remaining lineages also presented high support values for both
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FIGURE 1
A cladogram summarizing the phylogenetic relationships among Lasiodoriform genera and out-group lineages. Boxes on the nodes correspond to
the SH-aLRT test and Ultrafast Bootstrapping support values. Live specimen: Lasiodora sp. ♀. Photo credits: Wolf Moeller.
tests. Unfortunately, the placement of P. anomalus is still uncertain,
due to the low support value (bootstrapping = 86). The three
morphologically distinct taxa [Pterinopelma felipeleitei (Lasiodora
+ Pterinopelma sazimai)] were recovered as monophyletic, where
two of them would be considered monotypic genera: P. felipeleitei
and P. sazimai. Vitalius wacketi was recovered as the sister group
to Nhandu (Pterinopelma + Vitalius). Finally, the monophyletic
relationship of Pterinopelma and Vitalius is not well-supported,
with low SH-aLRT and bootstrap values (SH-aLRT = 80.5 and
bootstrapping = 82).
The genus Vitalius, as defined hitherto, was not recovered as
monophyletic (Figure 2). To recover the monophyly of the group, it
is necessary to transfer four species to other genera (see Taxonomy).
Within the genus Vitalius (sensu stricto), two clades were recovered
with high support values (SH-aLRT = 99.1 and bootstrapping =
90), the first comprising the type-species V. sorocabae sister group
of V. dubius (V. lucasae + V. buecherli) and the group with Vitalius
australis sp. nov. emerging as the sister group of V. paranaensis +
V. vellutinus.
Our results, in addition to not recovering the monophyly
of Vitalius, also showed that the genera Proshapalopus and
Pterinopelma, in their current conformation, are paraphyletic. To
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10.3389/fevo.2023.1177627
achieve taxonomic stability, we propose a series of changes in
the composition of the genera Pterinopelma and Vitalius, as well
as the description of three new genera and a new species (see
Taxonomy below). However, owing to the low support in the node
that joins Proshapalopus multicuspidatus and Vitalius nondescriptus
(see Figure 2), as well as the distant positioning of the type species
of the genus Proshapalopus anomalus, we chose to keep the genus
Proshapalopus paraphyletic.
3.2. Taxonomy
Taxonomic changes are proposed based on the resulting
phylogeny (Figures 1, 2) and morphological differences. New
morphological descriptions for all Lasiodoriform genera
(Figures 3–10) and the new species, including the taxonomic
changes, are found in Supplementary File S3. In this study, we
propose the following taxonomic acts: (1) transfer of Proshapalopus
amazonicus to Eupalaestrus; (2) transfer of Pterinopelma felipeleitei
to Parvicarina gen. nov. and of Pterinopelma sazimai to
Lasiocyano gen. nov.; and (3) transfer of Vitalius nondescriptus
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FIGURE 2
A phylogeny showing the relationships of Lasiodoriform genera. Boxes on the nodes correspond to the SH-aLRT test and Ultrafast Bootstrapping
support values. Lasiodoriform lineages are highlighted in pink.
to Proshapalopus, Vitalius wacketi to Tekoapora gen. nov., and
Vitalius roseus and Vitalius longisternalis to Pterinopelma.
Below, we have provided information for each genus belonging
to the Lasiodoriforms, in particular, type species, species included
in the genus, diagnoses and distribution, and a diagnosis for Vitalius
australis sp. nov.
Theraphosidae Thorell, 1869
Subfamily Theraphosinae
Eupalaestrus Pocock, 1901
Type-species: Eupalaestrus pugilator Pocock, 1901, by
original designation.
Contents: Eupalaestrus campestratus (Simon, 1891),
Eupalaestrus crassimetatarsis Borges et al., 2021, Eupalaestrus
larae Ferretti and Barneche, 2012, Eupalaestrus roccoi Borges
et al., 2021, Eupalaestrus spinosissimus Mello-Leitão, 1923 and
Eupalaestrus weijenberghi (Thorell, 1894), and Eupalaestrus
anomalus (Mello-Leitão, 1923) new comb., nomen rest.
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10.3389/fevo.2023.1177627
Emended diagnosis: Eupalaestrus males (except E. anomalus)
differ from those of all other theraphosine genera by the presence
of a denticulate row on the subapical area of the male palpal bulb
embolus (see Figures 50, 51 in Bertani, 2001) in conjunction with
the lack of stridulatory setae on the palpus trochanter. E. anomalus
males differ from other theraphosine genera by the thickened tibia
IV (see Figure 20 in Bertani, 2001). Eupalaestrus females differ
from those of all other theraphosine genera, except Vitalius lucasae,
by having the tibia IV thickened (Figure 3A), in conjunction with
scopula on the retrolateral face of femur IV and the absence of
stridulatory setae on the retrolateral face of the trochanter of the
palpus. They differ from V. lucasae by the presence of type III
urticating setae (except E. anomalus) or by the presence of stiff
setae on metatarsus IV (E. spinosissimus). Females of E. anomalus
differ from V. lucasae by their distinct geographic distribution (E.
anomalus occurs in the Amazon forest, while V. lucasae occurs in
the southern region of Brazil).
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FIGURE 3
Some Lasiodoriform live specimens. (A) Eupalaestrus campestratus ♀; (B) Eupalaestrus anomalus ♀; (C) Lasiodora sp. ♀; (D) Nhandu carapoensis ♀; (E)
Proshapalopus anomalus ♀; and (F) Vitalius sorocabae ♀. White arrows indicate the thickened tibia IV. Photo credits: (D) Wolf Moeller. (E) Rafael P.
Indicatti.

Distribution: Except for E. anomalus (from Brazilian Amazon,
state of Mato Grosso) and E. spinosissimus (Brazilian Atlantic
Forest, state of São Paulo and Rio de Janeiro), the genus is
predominantly found in open areas, occurring in Brazilian Cerrado
(state of Mato Grosso do Sul), Chaco (Argentina, Brazil, and
Paraguay), Pampa (Argentina, Brazil, and Uruguay), and Atlantic
Forest (Argentina, Brazil, and Paraguay).
Remarks: Eupalaestrus anomalus is restored into the
genus. Previously, Lucas et al. (1993) transferred this species
from Pocock, 1901 to Eupalaestrus anomalus. Later, Bertani
(2001) transferred the species to Proshapalopus based on the
morphological phylogenetic analysis, necessitating replacing the
name with Proshapalopus amazonicus. We transferred the species
to Eupalaestrus because it appears as a sister group to two other
species in our phylogeny. However, its distribution (Brazilian
Amazon) is far from the range of the other species of the genus
(southern Brazil, Paraguay, northern Argentina, and Uruguay).
Proshapalopus (Mello-Leitão, 1923)
Type species: Proshapalopus anomalus (Mello-Leitão, 1923)
Contents: P. anomalus, P. multicuspidatus (Mello-Leitão,
1929), and Proshapalopus nondescriptus (Mello-Leitão, 1926)
new comb.
Diagnosis: Males can be distinguished from other
Theraphosinae (except Eupalaestrus anomalus and Vitalius

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Galleti-Lima et al.
lucasae) by the presence of an accentuated male palpal bulb ventral
median depression (see Figures 68, 69 in Bertani, 2001; Figures 4B,
E). It differs from Eupalaestrus anomalus by the presence of
non-thickened tibia IV (Figure 3B), from Vitalius lucasae by the
presence of an accessory keel under the prolateral inferior keel
on the male palpal bulb (P. anomalus and P. multicuspidatus)
(see Figure 73 in Bertani, 2001; Figure 4E), bifid tibial spur with
very narrow branches, and the presence of long brown hairs
on metatarsus IV (P. nondescriptus) (see Figure 3 in Bertani
et al., 2012; Figure 4A). Females can be distinguished from other
Theraphosinae by the presence of type I urticating setae with
region “a” being shorter than region “b” (see Figure 9 in Bertani
et al., 2012).
Distribution: BRAZIL: Atlantic Forest, from extreme southeast
of the state of Minas Gerais to the states of Rio de Janeiro and
Espírito Santo, south of Rio Doce River (P. anomalus), Atlantic
Forest, from the state of Paraiba to south of the state of Bahia and
Chapada Diamantina (portion from Bahia) (P. multicuspidatus)
and from the state of Minas Gerais, in the highlands (up
1,100 m a.s.l.) of Espinhaço, Itacolomi, Caraça, and Gandarela (P.
nondescriptus) (Bertani, 2001; Bertani et al., 2012).
Remarks: Bertani et al. (2012) redescribed and transferred the
holotype Hapalopus nondescriptus Mello-Leitão, 1926 to Vitalius
nondescriptus (Mello-Leitão, 1926), based on the presence of a
subapical keel. Furthermore, the authors described the female
for the first time, emphasizing only the presence of type I
urticating setae, with region “a” being shorter than region “b”
(typical of Proshapalopus species). We decided to transfer this
species to the genus Proshapalopus based on our resulting
phylogenetic topology and morphological characteristics shared by
the species.
Parvicarina Galleti-Lima, Hamilton, Borges and Guadanucci,
new genus
ZooBank registration: urn:lsid:zoobank.org:act:D50EDC50-
41CC-4DFD-886F-2F0D2C3FD7CC.
Type species: Pterinopelma felipeleitei Bertani and Leal, 2016,
by monotypy.
Contents: Parvicarina felipeleitei (Bertani and Leal, 2016)
new comb.
Etymology: Generic name from the composition of the Parvi
(=small) and carina (=keel), in reference to the small keels present
on the male palpal bulb of the type-species. The genus name
is feminine.
Diagnosis: Males of Parvicarina felipeleitei can be distinguished
from other theraphosine genera by the following combination
of characters: weakly developed palpal bulb keels, almost
imperceptible (Figures 5C–F), and strongly curved metatarsus I
(see Figure 12 in Bertani and Leal, 2016) touching the apex of
the retrolateral spur when flexed (Figure 5G). Bifid tibial spur with
the apical region of the retrolateral branch curved prolaterally
(Figure 5E; see Figures 11, 12 in Bertani and Leal, 2016). Females
can be distinguished by the spermathecae separated by a sclerotized
area, which come up to half of the poorly sclerotized spermathecal
stalks (Figure 5B); absence of stridulatory setae on prolateral face of
coxa I; absence of long setae on the carapace (Figure 5A); absence
of type III urticating setae; sternum as long as wide (see Figure
9 in Bertani and Leal, 2016); and carapace and legs black with
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10.3389/fevo.2023.1177627
conspicuous white rings on distal femora, patellae, tibiae, and
metatarsi (Figure 5A; see Figures 20, 21 in Bertani and Leal, 2016).
Distribution: BRAZIL: Serra do Arrependido Condado
(Parque Estadual Pico do Itambé, municipality of Serro) and
PARNA Serra do Cipó, which includes the municipalities of
Morro do Pilar and Santana do Riacho. It also has records from
the municipalities of Gouveia and Diamantina. The sites where
individuals of the species were found belong to the Espinhaço
Mountain Range and are located in the central portion of the state
of Minas Gerais in Brazil (Bertani and Leal, 2016) (Figure 11A).
Lasiocyano Galleti-Lima, Hamilton, Borges, and Guadanucci,
new genus
ZooBank registration: urn:lsid:zoobank.org:act:653A6545-
E873-4071-B9EC-42146430C462.
Type species: Pterinopelma sazimai Bertani et al., 2011,
by monotypy.
Contents: Lasiocyano sazimai (Bertani et al., 2011) new comb.
Etymology: Generic name from the composition of Lasio
(=hairy) and Cyano (=blue), in reference to the blue iridescent
setae of the type-species. The genus name is neuter.
Diagnosis: Males of Lasiocyano sazimai can be distinguished
from all theraphosine genera by the palpal bulb with well-developed
prolateral superior, prolateral inferior, retrolateral, subapical, and
apical keels (Figures 6C, D; see Figures 6, 7 in Bertani and Leal,
2016); embolus slender at its distal portion (Figures 6C, D; see
Figures 6, 7 in Bertani and Leal, 2016); the absence of stridulatory
setae on prolateral face of coxa I; bifid tibial spur with apical region
of retrolateral branch curved prolaterally (Figure 6E; see Figure 5
in Bertani and Leal, 2016); and metatarsus I touching the apex
of retrolateral tibial spur when flexed (Figure 6E). Females of this
species can be distinguished by having the following combination
of traits: small spermatheca with spermathecal stalks proportional
to the size of the spermathecal receptacles (Figure 6B); the absence
of stridulatory setae on the prolateral face of coxa I; the absence
of long setae on the carapace (Figure 6A); and the presence of
type III urticating setae; Additionally, both males and females have
carapace, chelicerae, and legs covered with blue or purple iridescent
setae, which can vary to darker shades (Figure 6A; see Figures 1, 19
in Bertani and Leal, 2016).
Distribution: BRAZIL: Highlands (up 1,100 m a.s.l.) of the
Espinhaço Mountain range and States of Minas Gerais and Bahia,
Brazil (Bertani et al., 2011; Bertani and Leal, 2016) (Figure 11B).
Tekoapora Galleti-Lima, Hamilton, Borges, and Guadanucci,
new genus
ZooBank registration: urn:lsid:zoobank.org:act:BE01E016-
45A9-48A1-A0B2-7D80D04507BE.
Type species: Vitalius wacketi (Mello-Leitão, 1923),
by monotypy.
Contents: Tekoapora wacketi (Mello-Leitão, 1923) new comb.
Etymology: Generic name is derived from the way that the
original people from the Serra do Mar region (Mbyá) referred
to this geological formation (“tekoá porã”, means “good land”)
(Azanha and Ladeira, 1988). The genus name is masculine.
Diagnosis: Males differ from other theraphosine species by
having a palpal bulb with a long embolus, strongly pronounced
keels (Figures 7C, D), and male spurs with a prolateral branch,
which is smaller and more thickened than the retrolateral branch
07 frontiersin.org
Galleti-Lima et al. 10.3389/fevo.2023.1177627

FIGURE 4
“Proshapalopus” spp. (A) Proshapalopus nondescriptus ♂ live specimen. (B) Proshapalopus nondescriptus male palpal bulb, prolateral view. (C)
Proshapalopus nondescriptus male palpal bulb, retrolateral view. (D) Proshapalopus multicuspidatus (male juvenile) live specimen. (E) Proshapalopus
multicuspidatus male palpal bulb, prolateral view. (F) Proshapalopus multicuspidatus, retrolateral view. An arrow showing the dense tuft of brown
setae in the apical region of tarsus IV. A, Apical keel; AC, Accessory keel; D, Ventral median depression; PI, Prolateral inferior keel; PS, Prolateral
superior; R, Retrolateral keel; SA, Subapical keel. Scale bar: 1 mm. Photo credits: (A, C) Wolf Moeller.

(Figure 7E). Females can be distinguished by having tibiae IV
not thickened (Figure 7A); sternum slightly longer than wide;
carapace not bordered by long hairs pointing to the center carapace
(Figure 7A); big spermathecae with spermathecal receptacles not
enlarged (Figure 7B); femora with many long ventral setae; and
sternum and coxae black, covered by short hairs and the presence
of leg rings and longitudinal leg stripes being easily distinct.
Distribution: BRAZIL: Tropical latifoliated (broadleaf) forest
of “Serra do Mar” and coastal region, including many islands, from
south of the state of Rio de Janeiro and southwards to states of São
Paulo, Paraná, and Santa Catarina (Bertani, 2001) (Figure 11C).
Pterinopelma Pocock, 1901
Type-species: Pterinopelma vitiosum (Keyserling, 1891), by
original designation.
Contents: P. vitiosum, Pterinopelma longisternalis (Bertani,
2001) new comb. and Pterinopelma roseus (Mello-Leitão, 1923)
new comb.
Diagnosis: Pterinopelma shares with Lasiodora, Parvicarina
gen. nov., Vitalius, and Nhandu the absence of accessory prolateral
keels, as well as a ventral median region depression on the
palpal bulb and the presence of prolateral superior, prolateral
inferior, retrolateral, subapical and apical palpal keels in male bulb
(Figures 9A, B, D, E, G, H); Females differ from all other genera
in this study by the shape of the spermathecae with receptacles
separated by a small central depression and spermathecal stalk
with a slight constriction at the base of the spermathecal bulb
(Figures 8B, D, F). Both sexes can be distinguished from Lasiodora
by the absence of the stridulatory setae on the prolateral coxae.
Pterinopelma males are distinguished from Nhandu and Vitalius
by the palpal bulb shape that narrows abruptly from its median
region forward, a long embolus, denticles in the prolateral
inferior keel (P. vitiosum) (Figure 9A), and remarkable subapical
keel (P. longisternalis and P. roseus) (Figures 9D, E, G, H).
Females of Pterinopelma can be distinguished from Nhandu by
the absence of long setae on the carapace (Figures 8A, C, E)
and from Vitalius by having a spermathecal stalk narrower than
the spermathecal bulb, giving it a rounded shape (Figures 8B,
D, F).
Distribution: ARGENTINA: Araucaria forests northeastern of
Misiones province (P. longisternalis) and southeastern of Misiones
province (P. roseus). BRAZIL: northern of the state of Rio Grande
do Sul, where vegetation comprises Subcaducifolious Atlantic
Forest with Araucaria angustifolia and patches of “campo” (P.
vitiosum), subtropical subcaducifolious forest of states of Paraná
and Santa Catarina, west of “Serra do Mar” (P. longisternalis), and
subtropical subcaducifolious forest of northwest and central state

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FIGURE 5
Parvicarina felipeleitei. (A) Parvicarina felipeleitei ♀ live specimen. (B) Ventral face of spermathecae. (C) Parvicarina felipeleitei male palpal bulb,
prolateral view. (D) Prolateral view of the embolus in detail. (E) Parvicarina felipeleitei male palpal bulb, retrolateral view. (F) A retrolateral view of the
embolus in detail. (G) Parvicarina felipeleitei tibial apophysis and flexion of the metatarsal I on the tibial apophysis. A, Apical keel; PI, Prolateral inferior
keel; PS, Prolateral superior; R, Retrolateral keel. Scale bar: (A–G) = 1 mm; (D, F) = 0.5 mm. Photo credits: (A) Pedro H. Martins.

of Rio Grande do Sui (P. roseus) (Bertani, 2001; Bertani et al., 2011;
Ferretti et al., 2014, 2015) (Figure 11D).
Vitalius Lucas et al., 1993
Type-species: Vitalius sorocabae (Mello-Leitão, 1923), by
original designation.
Contents: V. sorocabae, V. buecherli, V. dubius, V. lucasae, V.
paranaensis, V. vellutinus, and Vitalius australis new species
Emended diagnosis: Males can be distinguished from species
of other theraphosine genera except Proshapalopus, Lasiodora,
Lasiocyano gen. nov., Tekoapora gen. nov., Nhandu, and
Pterinopelma by the presence of a triangular subapical keel
on the male palpal bulb (see Figures 74–79, 91–96, 101–104,
119, 120, 123, 124, 127, 128 in Bertani, 2001; Figures 10C, D).
They can be distinguished, from Lasiodora, by the absence of
a velvet stridulating setae on the coxae; from P. anomalus and
P. multicuspidatus, by the absence of an acessory keel in the
male palpal bulb; from P. nondescriptus, except for V. lucasae,
by the absence of accentuated ventral median depression on the
palpal bulb; from Nhandu, except for V. lucasae, by the male
spur with converging branches originating from a common base,
tapering distally, with the prolateral branch thickened or by small,
almost vestigial, male spurs (V. vellutinus) (see Figures 108–111
in Bertani, 2001). Moreover, V. lucasae can be distinguished from
P. nondescriptus and Nhandu by the thickened tibiae IV. Males
of Vitalius differ from Lasiocyano gen. nov. by the absence of
blue color patter and the apical region of retrolateral branch not
curved prolaterally (see Figures 80, 97, 108–111, 117, 121, 125
in Bertani, 2001; Figure 10E); from Tekoapora gen. nov. by the
not thickened both tibial aphophysis branches (see Figures 80, 97,
108–111, 117, 121, 125 in Bertani, 2001; Figure 10E); and from
Pterinopelma by the palpal bulb shape that not narrows abruptly
from its median region forward (see Figures 74–79, 91–96, 101–
104, 119, 120, 123, 124, 127, 128 in Bertani, 2001; Figures 10C, D).
Females can be distinguished from Lasiodora by the absence of a
velvet stridulating setae on the coxae, from Proshapalopus by the
Type I urticating setae with region “A” longer or equal to region
“B”; from Lasiocyano gen. nov. by the absence of blue color and
Type III urticating setae; from Nhandu by the absence of many
long, curly, scattered hairs over the carapace; from Tekoapora gen.
nov. by the small spermathecae; and from all Pterinopelma species
by the spermathecae without a small central depression separating
the receptacles (see Figures 81, 82, 98–100, 107, 118, 122, 126 in
Bertani, 2001; Figure 10B).
Distribution: ARGENTINA: Misiones province; BRAZIL:
States of São Paulo, Paraná, and Rio Grande do Sul (Figures 11E,
F).
Vitalius australis Galleti-Lima, Hamilton, Borges, and
Guadanucci, new species
ZooBank registration: urn:lsid:zoobank.org:act:57A4852C-
84CA-425E-94DF-340656BE3142.

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Galleti-Lima et al. 10.3389/fevo.2023.1177627

FIGURE 6
Lasiocyano sazimai. (A) Lasiocyano sazimai ♀ live specimen. (B) Ventral face of spermathecae. (C) Lasiocyano sazimai male palpal bulb, prolateral
view. (D) Lasiocyano sazimai male palpal bulb, retrolateral view. (E) Lasiocyano sazimai tibial apophysis and flexion of the metatarsal I on the tibial
apophysis. A, Apical keel; PI, Prolateral inferior keel; PS, Prolateral superior; R, Retrolateral keel; SA, Subapical keel. Scale bar: 1 mm. Photo credits: (A)
Wolf Moeller.

Type material: Holotype: (CAD 1244). Paratypes: (CAD 1210, 4. Discussion

CAD 1243).
Etymology: Generic name in reference to the region where
the species is found (Southern Region of Brazil). The genus name
is male.
Diagnosis: Males can be distinguished from congeners by the
long embolus, very pronounced prolateral superior and inferior
keels, and a well-developed male spur, with the retrolateral
branch tapering distally (Figures 10C, D). Females differ from
the other species by the short and homogeneously sclerotized
spermatheca, with a spermathecal stalk with a strong constriction
at the base of the spermathecal bulb that does not have
a rounded shape, and also by the elevated central region
(Figure 10B).
Distribution: Probably from the Seasonal Deciduous
Forest in the north of Rio Grande do Sul to the
Araucaria Forest region in the interior of Paraná
(Figure 11E).
4.1. Lasiodoriforms: theraphosinae with
subapical and retrolateral keels
The presence of keels on the male palpal bulb is a
synapomorphic character of Theraphosinae (Pérez-Miles et al.,
1996). These keels were first proposed to have taxonomic value
for tarantulas by Bücherl (1957) and later, Bertani (2000) proposed
terminology, homologies, and detailed descriptions of palpal
bulb keels of Theraphosinae genera, regarding their morphology,
position, and distribution among this genera. In a morphological,
phylogenetic, and biogeographic study of the genera Vitalius,
Nhandu, and Proshapalopus, Bertani (2001) suggested the presence
of a retrolateral keel and a male palpal bulb with the embolus
apex slightly laterally flattened, characterized by concave areas
above and under this retrolateral keel, as synapomorphies of
Eupalaestrus, Lasiodora, Proshapalopus, Nhandu, and Vitalius

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FIGURE 7
Tekoapora wacketi. (A) Tekoapora wacketi ♀ live specimen. (B) Ventral face of spermathecae. (C) Tekoapora wacketi male palpal bulb, prolateral view.
(D) Tekoapora wacketi male palpal bulb, retrolateral view. (E) Tekoapora wacketi tibial apophysis and flexion of the metatarsal I on the tibial
apophysis. A, Apical keel; PI, Prolateral inferior keel; PS, Prolateral superior; R, Retrolateral keel; SA, Subapical keel. Scale bar: 1 mm. Photo credits: (A)
Rafael P. Indicatti.

clades. Bertani (2001) also found that Lasiodora, Proshapalopus,
Nhandu, and Vitalius formed a monophyletic group supported
by the presence of a subapical keel and males with metatarsus I
that touches the retrolateral branch of the tibial apophysis when
flexed (except in Vitalius species), and the genus Eupalaestrus, as
sister-group. Our analysis supports the monophyly of the group,
and we propose the name Lasiodoriform to include: Eupalaestrus,
“Proshapalopus”, Parvicarina gen. nov., Lasiodora, Lasiocyano gen.
nov., Tekoapora gen. nov., Nhandu, Pterinopelma, and Vitalius.
The genus Lasiodora is the oldest described genus among the taxa
of this group, and all the genera share sexual characteristics similar
to Lasiodora, such as bulbs with subapical and retrolateral keels
(except for the male bulbs of some species of Eupalaestrus and
Parvicarina gen. nov.), male tibial apophysis with two branches
(except for some species of Nhandu, who had a secondary loss of the
tibial aphophysis), and spermathecae fused in a small central area.
51 in Bertani, 2001), which is found in the same position as
the subapical keel but is morphologically distinct. Bertani (2000,
2001) suggested that the denticulate row of Eupalaestrus is a
homolog feature to the subapical keel of Lasiodora, Proshapalopus,
Nhandu, and Vitalius. In this study, we assume that the genus
Eupalaestrus belongs in the Lasiodoriform lineage. Eupalaestrus
anomalus (Figure 3B), which emerged as a sister lineage to the other
species of the genus, has a palpal bulb with a small and triangular
subapical keel, which is almost inconspicuous (see Figures 64,
65 in Bertani, 2001). This finding putatively corroborates that
the denticulate row of Eupalaestrus is a primary homolog of the
subapical keel, as proposed by Bertani (2000, 2001), making this
characteristic a synapomorphy of the genus that is absent in E.
anomalus. All species of Eupalaestrus have males and females with
thickened tibia IV, making this an important synapomorphy of
the genus.

4.3. The non-monophyly of Proshapalopus

4.2. The genus Eupalaestrus and the species
E. anomalus
The monophyly of the genus Eupalaestrus (Figures 3A, B) is
supported by the presence of a denticulate row (see Figures 50,
Proshapalopus multicuspidatus is related to P. nondescriptus,
previously included in Vitalius. P. nondescriptus was previously
included in Vitalius because females lack type III urticating setae
and males have the palpal bulb with a well-pronounced subapical

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FIGURE 8
Pterinopelma. (A) Pterinopelma vitiosum ♀ live specimen. (B) The ventral face of spermathecae of P. vitiosum. (C) Pterinopelma longisternalis ♀ live
specimen. (D) The ventral face of spermathecae of P. longisternalis. (E) Pterinopelma roseus ♀ live specimen. (F) Ventral face of spermathecae of P.
roseus. Scale bar: 1 mm. Photo credits: (C) Rafael P. Indicatti.

keel (Bertani et al., 2012). Here, we present morphological
characteristics that support the relationship between P.
multicuspidatus and P. nondescriptus, such as the presence of
a ventral median depression (D) on a male palpal bulb (Figures 4B,
C, E, F) and females with type I urticating setae with region “a”
shorter than region “b” (see Figure 3 in Bertani, 2001 and Figure
9 in Bertani et al., 2012). Females of P. anomalus also present this
same variation of type I urticating setae, in addition to males with
palpal bulbs possessing an accessory keel (AC), a character shared
with males of P. multicuspidatus. One potential issue is that the
relation of P. multicuspidatus + P. nondescriptus has low support
in both tests (SH-aLRT and bootstrapping). We expected to find
P. anomalus, P. multicuspidatus and P. nondescriptus in the same
clade because they share these characteristics; however, we did not
recover these species as monophyletic in our topology. To restore
the monophyly of Vitalius (the focus of this study), we propose here
the transfer P. nondescriptus to Proshapalopus, which now includes
P. anomalus, P. multicuspidatus, and P. nondescriptus, keeping the
genus Proshapalopus paraphyletic (see topology in Figures 1, 2).
We presumed that the relationship between P. multicuspidatus, P.

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FIGURE 9
Male sex characteristics of Pterinopelma species. (A) P. vitiosum male palpal bulb, prolateral view. (B) P. vitiosum male palpal bulb, retrolateral view.
(C) P. vitiosum tibial apophysis and flexion of the metatarsal I on the tibial apophysis. (D) P. longisternalis male palpal bulb, prolateral view. (E) P.
longisternalis male palpal bulb, retrolateral view. (F) P. longisternalis tibial apophysis and flexion of the metatarsal I on the tibial apophysis. (G) P.
roseus male palpal bulb, prolateral view. (H) P. roseus male palpal bulb, retrolateral view. (I) P. roseus tibial apophysis and flexion of the metatarsal I on
the tibial apophysis. A, Apical keel; PI, Prolateral inferior keel; PS, Prolateral superior; R, Retrolateral keel; SA, Subapical keel. Scale bar: 1 mm.

nodescriptus, and P. anomalus can be recovered with additional
analyses that include more representatives of these taxa.
We must highlight the absence of the recently described
genus Cymbiapophysa (Gabriel and Sherwood, 2020), from
Ecuador, Colombia, and possibly Peru (Gabriel and Sherwood,
2020; Sherwood et al., 2021). The genus was described based
on the holotype of Metriopelma velox Pocock, 1903 and
comprises four species, including Cymbiapophysa marimbai
(Perafán and Valencia-Cuellar, 2018), which was previously
described in the genus Proshapalopus, based on its position in
a morphological phylogeny, and supported by morphological
characteristic diagnostic of Proshapalopus (Perafán and Valencia-
Cuellar, 2018). It is worth noting that the authors emphasized
the disruptive distribution of this taxa in comparison to
other species of Proshapalopus and discussed the hypothesis
with which this taxon could form a new taxonomic group.

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FIGURE 10
Vitalius australis sp. nov. (A) Vitalius australis sp. nov. ♀ live specimen. (B) Ventral face of spermathecae. (C) Vitalius australis sp. nov. male palpal bulb,
prolateral view. (D) Vitalius australis sp. nov. male palpal bulb, retrolateral view. (E) Vitalius australis sp. nov. tibial apophysis and flexion of the
metatarsal I on the tibial apophysis. A, Apical keel; PI, Prolateral inferior keel; PS, Prolateral superior; R, Retrolateral keel; SA, Subapical keel. Scale bar:
1 mm. Photo credits: (A) Wolf Moeller.

Based on the morphology of the male palpal bulb, tibial
apophysis, and female spermathecae, Sherwood et al. (2021)
transferred this species to Cymbiapophysa and questioned the
designation of the keels and palpal bulb structures of the
species by Perafán and Valencia-Cuellar (2018), proposing a
new terminology. The support to include this species in
Cymbiapophysa is weak, given that C. marimbai lacks cymbial
apophysis (found in other species of the genus), presenting
only a sclerotized portion of cymbium, as noted by Perafán
and Valencia-Cuellar (2018), but no phylogeny was presented to
test the monophyly of the genus. Sherwood et al. (2021) also
observed a similar cymbial apophysis of other Cymbiapophysa
species on P. amazonicus (here transferred to Eupalaestrus).
We were unable to insert any Cymbiapophysa (including C.
marimbai) in our analysis to clarify its phylogenetic position.
However, we agree with the hypothesis presented by Perafán
and Valencia-Cuellar (2018) that C. marimbai may be related
to an undescribed lineage, but new taxonomic and phylogenetic
studies would be necessary to test the hypothesis and clear the
genus positioning.
4.4. Clade formed by Parvicarina gen. nov.,
Lasiodora, and Lasiocyano gen. nov.
The monophyly of this group (Figures 3C, 5A, 6A) is well-
supported in the phylogenomic analysis and can be further
supported by the morphology of the male tibial apophysis with
the apical region of the retrolateral branch curved prolaterally
(Figures 5E, 6E; see Figure 58 in Bertani, 2001). Considering the
topology of the tree, two options are possible: (1) if the whole clade
can be considered Lasiodora, then the diagnoses of the genus should
be amended to include two extra species; or (2) keep Lasiodora as it
is, diagnosed by the presence of a stridulating setae on coxae I and
II above the suture, and propose two new monotypic genera. The
synonymy between these genera was not considered here because
the three taxa have other distinct and exclusive characters, those of
which warrant generic recognition (see diagnoses in Taxonomy).
Because of this, we chose the second option, a decision that agrees
with the repeatedly supported monophyly of Lasiodora (Bertani,
2001; Bertani et al., 2011; Galleti-Lima and Guadanucci, 2018;
Perafán and Valencia-Cuellar, 2018). Moreover, the two novel

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FIGURE 11
A distribution map of rearranged Lasiodoriform species according to the topology. (A) Parvicarina felipeleitei. (B) Lasiocyano sazimai. (C) Takoapora
wacketi. (D) Pterinopelma spp. (E) V. paranaensis, V. vellutinus, and Vitalius australis sp. nov. (F) V. buecherli, V. dubius. V. lucasae, and V. sorocabae.

genera can be diagnosed by the absence of a subapical keel, as well According to records in the literature (Bertani et al.,
as poorly-developed keels for Parvicarina gen. nov. and by the body 2011; Bertani and Leal, 2016; Galleti-Lima and Guadanucci,
color with iridescent blue setae on the carapace, chelicerae, and legs 2018) and scientific collections, Lasiocyano sazimai has exclusive
in Lasiocyano gen. nov. distribution of rocky fields of the Espinhaço Mountain Range

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Galleti-Lima et al.
of the states of Bahia and Minas Gerais, a region that has
been heavily impacted by anthropogenic action over the years
(ICMBio, 2018). Another important factor to note is the
illegal trade in wild animals (ICMBio, 2018), which, along
with habitat loss, has resulted in the Lasiocyano sazimai being
listed as endangered on the official list of Brazilian terrestrial
invertebrate animals threatened with extinction (see list in Leite,
2021).
4.5. Clade formed by Tekoapora gen. nov.,
Nhandu, Pterinopelma, and Vitalius
All past morphological phylogenies that focused on
Lasiodoriforms have recovered the sister group relation Nhandu
+ Vitalius (Bertani, 2001; Bertani et al., 2011; Galleti-Lima and
Guadanucci, 2018; Perafán and Valencia-Cuellar, 2018). Although
the revalidation of Pterinopelma was based on phylogenetic
inference, two of the three species (P. sazimai and P. felipeleitei)
were not analyzed in a phylogenetic context after misidentifications
were pointed out (Bertani and Leal, 2016). A recent morphology-
based phylogenetic analysis that included Pterinopelma species
(Galleti-Lima and Guadanucci, 2018) challenged the monophyly
of Nhandu + Vitalius, where P. sazimai appeared as a sister group
to the genus, Nhandu. Females of P. vitiosum share morphological
similarities with these other two genera, especially Vitalius, due
to the absence of type III urticating setae in females and their
color pattern. These similarities are so evident that P. longisternalis
and P. roseus were placed in Vitalius by Bertani (2001), and
only with the use of phylogenomics was it possible to verify the
proximity of these two species to P. vitiosum. We observed high
morphological similarity between Vitalius, and the species included
herein is Pterinopelma (P. roseus, P. longisternalis, and P. vitiosum).
Owing to these findings, we propose a set of morphological
characters distinguishing Vitalius from Pterinopelma and other
Lasiodoriforms (see diagnosis for these genera in Taxonomy).
The genera that constitute the Tekoapora gen. nov., Nhandu,
Pterinopelma, and Vitalius clade have subtle morphological
diagnoses that are difficult to be recognized. Despite this,
we decided to keep the three genera due to the support of
the nodes forming monophyletic groups and the placement
of the type species in each of these groups. Tekoapora gen.
nov., a sister-group of the other genera, has a combination
of characteristics that appeared multiple times and individually
along the other related genera, such as the presence of a
long embolus maintained only in Pterinopelma or the strongly
pronounced keels in the male palpal bulb, observed in Nhandu
but not in other genera. Furthermore, Tekoapora gen. nov.
appears to be endemic to the Serra do Mar, a mountain
system that extends from the coastal region of Espírito Santo
to the south of Santa Catarina, and which was raised by
the tectonic action during the post-Cretaceous (Ross, 1996;
Bertani, 2001). Bertani (2001) suggested that this geological
event could be related to the distribution of Vitalius species
(extrapolating here to all genera of this clade, since the vast majority
are sympatric).
Frontiers in Ecology and Evolution
10.3389/fevo.2023.1177627
4.6. The intrarelationships of Vitalius
Among the Vitalius species, there are two recognizable groups:
one formed by V. buecherli, V. lucasae, V. dubius, and V. sorocabae
and another formed by V. paranaensis, V. vellutinus, and Vitalius
australis sp. nov. The proximity between the species V. dubius and
V. buecherli has been discussed by Bertani (2001), who pointed
to the similarity of the palpal bulb of these species (see Figures
91–96, 123, 124 in Bertani, 2001). Furthermore, the position of
V. lucasae as the sister lineage to V. buecherli differs from the
morphological phylogenetic hypotheses, where the species was the
sister group to all other Vitalius species. The distribution of these
species appears to be geographically continuous, extending through
the Atlantic Plateau, with a broadleaf tropical forest vegetation.
This was suggested by Bertani (2001) as a factor of sympatry
between V. dubius and V. buecherli. Additionally, Bertani (2001)
suggested that the areas of occurrence of V. dubius and V. buecherli
were also interconnected with V. lucasae through the distribution
area of V. sorocabae, which here emerged as a sister group to all
these species.
Within the other group (V. paranaensis, V. vellutinus and
Vitalius australis sp. nov.), V. paranaensis is sister to V. vellutinus,
a relationship similar to what was discovered by Bertani (2001), but
without the proposition of well-defined morphological diagnostic
characters to distinguish between females. Bertani (2001) proposed
that females of these two taxa could only be recognized by
geographic distribution, with V. paranaensis restricted to the state
of Paraná and the province of Misiones, Argentina, while V.
vellutinus is distributed across the west of the state of São Paulo.
We recovered the relationship of two individuals identified as V.
vellutinus, one collected in the state of São Paulo and the other in
the state of Paraná, highlighting the need for further investigation
toward new diagnostic delimitation for these females. In addition,
the males of the new species (Vitalius australis sp. nov.) present
morphological characteristics similar to the males of V. paranaensis
and can only be differentiated due to their phylogenetic relationship
with a morphologically distinct female individual. From this, it was
possible to determine small morphological differences in the tibial
apophyses and male palpal bulb.
Data availability statement
The original contributions presented in the study are
publicly available. This data can be found here: ZooBank,
urn:lsid:zoobank.org:act:653A6545-E873-4071-B9EC-42146430
C462, urn:lsid:zoobank.org:act:D50EDC50-41CC-4DFD-886F-
2F0D2C3FD7CC, urn:lsid:zoobank.org:act:BE01E016-45A9-48A1-
A0B2-7D80D04507BE, and urn:lsid:zoobank.org:act:57A4852C-
84CA-425E-94DF-340656BE3142.
Ethics statement
Ethical review and approval was not required for the
study on animals in accordance with the local legislation and
institutional requirements.
16 frontiersin.org
Galleti-Lima et al.
Author contributions
AG-L and JG conceived the research that resulted in this
paper. CH developed and funded all molecular analyses, in
addition to collaborating on the manuscript. LB assisted with
specimen collection, taxonomic and morphological discussions,
and collaborated with the manuscript. The text was revised by all
authors. All authors contributed to the article and approved the
submitted version.
Funding
This research was funded by the Conselho Nacional de
Desenvolvimento Científico e Tecnológico—CNPq (Process No.
168493/2017-9) for AG-L, Fundação de Amparo à Pesquisa
do Estado de São Paulo—FAPESP (Process No. 2017/11985-9)
for JG, Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior—CAPES (Process No. 88881.709394/2022-01), (Process
No. 88882.428068/2019-01) for LB, and University of Idaho to CH.
Acknowledgments
We would like to express our gratitude to all curators
who provided access to the materials studied in this research.
We thank the technician (Andrea Noble Stuen) and graduate
students (Karina Silvestre Bringas and Erik Ciaccio) from the
Department of Entomology, Plant Pathology and Nematology in
the University of Idaho who were responsible for the sample
extractions. We are also grateful to the Instituto Chico Mendes
de Conservação da Biodiversidade—ICMBio (64478-1 and 68145-
1) and the Comissão Técnica Científica do Instituto Florestal—
COTEC (260108-004.071/2018), for granting authorizations for
specimen collection. We thank Pró-Reitoria de Pesquisa of Unesp
for the financial support to publish this article (Edital 01/2023
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 Supplementary File S3
Theraphosidae Thorell, 1869
Subfamily Theraphosinae
Eupalaestrus Pocock, 1901
Eupalaestrus anomalus (Mello-Leitão, 1923) new comb., nomen rest.
Mello-Leitão, 1923: 343 (male lectotype, not examined, and three males paralectotypes
from Santo Christo, Rio Tapajós, Pará, Brazil, MZSP 555, Garbe coll., not examined).
Eupalaestrus anomalus; Lucas, Silva & Bertani, 1993: 245. Proshapalopus
amazonicus; Bertani, 2001: 287, figs. 62-65, 175 (replacement name for P.
amazonicus Mello-Leitão, 1923, preoccupied). P. amazonicus; Peters, 2003: 272, figs.
1087, 1090-1091. P. amazonicus; Schmidt, 2003: 184, figs. 474-476. P.
amazonicus; Schmidt, 2007: 100, fig. 5. P. amazonicus; Sherwood et al., 2021: 839,
figs. 11-17.
Diagnosis: Males and females resembles other Eupalaestrus species by the thickened
tibia IV (Figures 3A-B). Males differ from other Eupalaestrus species by the absence of
a denticulate row on palpal bulb and by the small, triangular subapical keel bordered by
small denticles and a palpal bulb with accentuated D region (See Bertani, 2001, figs. 64-
65). Females can also be distinguished by the absence of type III urticating setae.
Description: See Bertani (2001).
Additional material examined: BRAZIL, Pará, Aveiro, Sítio Paraíso, 4°04'15.6"S,
55°26'49.2"W, 2 juveniles, 4-7.X.2020, R. Fonseca-Ferreira coll. (CAD 1155), female,
4-7.X.2020, R. Fonseca-Ferreira coll. (CAD 1156).

Proshapalopus Mello-Leitão, 1923

Description: Carapace longer than wide. Cephalic region slightly raised, convex.
Thoracic striae moderately deep. Fovea short, straight and shallow (P. nondescriptus) or
deep (P. multicuspidatus). Chelicerae without rastellum, basal segments with 10 to 12
teeth. Eye tubercle distinct, wider than long. Short clypeus. Anterior eye row procurved,
posterior recurved. Labium slightly wider than long, with numerous (100 to 200)
cuspules. Maxilla with numerous cuspules (100 to 200). Sternum longer than wide. First
pair of sigilla rounded, second and third oval. Leg formula: IV I II III. Clavate
trichobothria on distal 2/3 of tarsi I–IV. Absense of stridulatory apparatus. Femur IV
with retrolateral scopula. Leg tarsi without spines, claw tufts present. Tarsi I-IV and
metatarsi I-II scopulated, metatarsus III 1/3 scopulated (P. nondescriptus) or along half
its length (P. anomalus and P. multicuspidatus), metatarsus IV scopulate (P.
nondescriptus) or apically scopulated (P. anomalus and P. multicuspidatus). Type I and
III urticating hair present in males. Type I and III urticating hair present in females (P.
anomalus and P. multicuspidatus) or type III urticating hair absent in females (P.
nondescriptus). Type I urticating hair with the region "a" shorter than the region "b".
Male palpal bulb pyriform, embolus slightly flattened distally. Embolus short. Prolateral
keels present, the Prolateral Superior keel forming the embolus edge distally. Acessory
keel present, under of the Prolateral Inferior keel (P. anomalus and P. multicuspidatus).
Retrolateral keel present, not pronounced. Apical keel small. Subapical keel small,
triangular, bordered by small denticles. Male palpal bulb with accentuated (P.

1
multicuspidatus) or slightly (P. anomalus) ventral median depression. Male spur with
straight branches originating from a common base, the retrolateral constricted in the
middle (P. anomalus) or branches tapering distally, the prolateral straight and
retrolateral longest with slight distal curvature (P. nondescriptus) or male spur with
prolateral branche thickened (P. multicuspidatus). Metatarsus I straight, when flexed
touches the retrolateral branch of the male spur laterally (P. multicuspidatus and P.
nondescriptus) or the apex of the retrolateral branch (P. anomalus). Spermathecae short,
separated by a heavily (P. nondescriptus) or weakly (P. anomalus and P.
multicuspidatus) sclerotized area. Spermathecal stalk narrower than spermathecal bulb.
Carapace covered only by short slender setae, bordered by numerous short setae
pointing out. Slender legs with metatarsus IV covered by long brown setae (New Genus
1 nondescriptus). Coxae and sternum covered by short setae. Color pattern ranging from
Light brown, with femora slightly darker (P. anomalus), to homogeneous, dark gray,
with light brown setae on its border of carapace (P. nondescriptus) and to dark-brown
(P. multicuspidatus). Coxae, labium, sternum, maxila and legs orange (P. anomalus)
brown (P. nondescriptus) or black (New Genus 1 multicuspidatus). Leg rings and stripes
inconspicuous (New Genus 1 nondescriptus) or easly distinct (P. anomalus and P.
multicuspidatus). Abdomen dorsally black (P. nondescriptus) or brown (P. anomalus
and P. multicuspidatus).

Proshapalopus anomalus Mello-Leitão, 1923

Proshapalopus anomalus Mello-Leitão, 1923: 146, Figs. 60-62, 186; 190 (holotype
male from Pinheiro (= Pinheiral), Rio de Janeiro, Brazil, in MNRJ, not examined).
Petrunkevitch, 1928: 79; 1939: 281. Roewer, 1942: 236. Bonnet, 1958: 3780.
Stichoplastus anomalus; Raven, 1985: 157. Platnick, 1989: 110. Proshapalopus
anomalus; Rudloff, 1997: 2. P. anomalus; Bertani, 2001: 289, figs. 70-73, 177. P.
anomalus; Schmidt, 2003: 131, figs. 152-154.
Diagnosis: Males can be distinguished from other species by the absence of an
accentuated ventral median depression (See Bertani, 2001, figs. 72-73). They can also
be distinguished from P. multicuspidatus and P. nondescriptus by the metatarsus I
touching the apex of the retrolateral spur branch when flexed (See Bertani, 2001, fig.
70). Females can be distinguished from P. nondescriptus by the presence of urticating
setae type III and from P multicuspidatus, by the distinct color pattern and geographical
distribution.
Description: See Bertani (2001).
Additional material examined: BRAZIL, Espírito Santo: Santa Tereza, male,
08.X.2015 (CAD 573), female, V.2015, J. P. Simor coll. (CAD 1157), female, V.2015,
J. P. Simor coll. (CAD 1159); Rio de Janeiro: Itatiaia, Parque Nacional do Itatiaia, male,
XI.2016, V. M. Ghirotto coll. (CAD 1160), male, III.2019, A. Galleti-Lima, I. Meireles,
R. Fonseca-Ferreira & R. P. Indicatti coll. (CAD).

P. multicuspidatus (Mello-Leitão, 1929)

Phormictopus multicuspidatus Mello-Leitão, 1929: 91, Figs. 1-2 (holotype male from
Tapera, Pernambuco, Brazil, in MNRJ, Bento Pickel col., not examined). Cyclosternum
multicuspidatum; Bücherl, Costa & Lucas, 1971: 123, figs 21-24. Proshapalopus

2
multicuspidatus; Bertani, 2001: 288, figs. 24, 66-69, 176. P. multicuspidatus; Schmidt,
2003: 185, figs. 477-479.
Diagnosis: Males differ from P. nondescriptus by the presence of an accessory keel
below the prolateral inferior on the male palpal bulb (Figure 4F) and by the absense of a
dense tuft of brown setae in the apical region of metatarsus IV (Figure 4D). They can be
distinguished from P. anomalus by the metatarsus I touching the retrolateral spur branch
laterally when flexed and by the presence of an accentuated D (Fig. 4E-F). Females can
be distinguished from P. nondescriptus by the presence of urticating setae type III and
from P. anomalus by the color pattern and distinct geographical distribution.
Description: See Bertani (2001).
Additional material examined: BRAZIL, Alagoas: São Miguel dos Campos,
9°47'17.7"S, 36°06'00.1"W, male, 2018, A. D. Abegg coll. (CAD); Quebrangulo,
Reserva Biológica de Pedra Talhada, male, 26-27.IV.2022, M.D.F. Magalhães, P.H.
Martins & H.M.O. Gonzalez-Filho coll., (CAD); Bahia: São Desidério, male, VII.2014,
V. M. Ghirotto coll. (CAD 1158); Catolés, Mata da Tijuquinha, 13º16'09.23''S,
41º54'40.65''W, male, 02.III.2022, M. D. F. Magalhães, A. Galleti-Lima & P. H.
Martins coll (UFMG); Jussari, RPPN Serra do Teimoso - Pau Brasil trail,
15º09'17.83''S, 39º31'52.31''W, juvenile, 10.III.2022, M. D. F. Magalhães, A. Galleti-
Lima & P. H. Martins coll (UFMG); Una, Rebio Una, Main road, 15º11'26.97''S,
39º03'47.24''W, juvenile, 11.III.2022, M. D. F. Magalhães, A. Galleti-Lima & P. H.
Martins coll (UFMG). Paraíba: João Pessoal, Parque Estadual Mata do Buraquinho,
male and female, 18-19.IV.2022, M.D.F. Magalhães, P.H. Martins & H.M.O. Gonzalez-
Filho (CAD).

P. nondescriptus (Mello-Leitão, 1926) new comb.

Hapalopus nondescriptus Mello-Leitão, 1926: 319, figs. 7-8; Silva-Moreira et al., 2010:
69; Cyclosternum melloleitaoi Bücherl, Timotheo & Lucas, 1971: 125, figs. 29-31; First
synonymized by Silva-Moreira et al., 2010: 69; Vitalius nondescriptus; Bertani,
Nagahama & Fukushima, 2012: 468, figs. 1-11 (comb. n.) (Redescribed, Holotype
male, MNRJ 43, not examined).
Diagnosis: Males can be distinguished from P. multicuspidatus by the absence of an
accessory keel under the prolateral inferior on male palpal bulb (Figure 4B-C) and by
the presence of a dense tuft of brown setae in the apical region of tarsus IV (Fig. 4A).
They can be distinguished from P. anomalus by metatarsus I the touching the
retrolateral spur branch laterally when flexed. Females differ from other species by the
absence of urticating setae type III.
Description: See Bertani et al. (2012).
Additional material examined: BRAZIL, Minas Gerais: Diamantina, female (CAD
06), male, IX.2008, J. P. L. Guadanucci coll. (CAD 83), female, Zoonozes coll. (CAD
93), male, XI.2010, male, XI.2010 (CAD 442), Vila Operária, male, 01.XI.2011 (CAD
461), male, IX.2011, J. P. L. Guadanucci coll. (CAD 462), male, J. P. L. Guadanucci
coll. (CAD 464), male, 01.XI.2011, J. P. L. Guadanucci coll. (CAD 465), male,
X.2011, J. P. L. Guadanucci coll. (CAD 467), female (CAD 471), female (CAD 569),
female, 25.X.2018, A. Galleti-Lima, V. M. Ghirotto & R. Fonseca-Ferreira coll. (CAD
1203), female, 25.X.2018, A. Galleti-Lima, V. M. Ghirotto & R. Fonseca-Ferreira coll.

3
(CAD 1204), juvenile, 25.X.2018, A. Galleti-Lima, V. M. Ghirotto & R. Fonseca-
Ferreira coll. (CAD 1205), female, 25.X.2018, A. Galleti-Lima, V. M. Ghirotto & R.
Fonseca-Ferreira coll. (CAD 1206), juvenile, 25.X.2018, A. Galleti-Lima, V. M.
Ghirotto & R. Fonseca-Ferreira coll. (CAD 1207); São Gonçalo do Rio da Pedra, male,
19-25.X.2010, P. Louise, J. P. L. Guadanucci & W. F. Silva coll. (CAD 498); Santana
do Riacho, Parque Nacional da Serra do Cipó, male, 17-20.X.2018, A. Galleti-Lima, V.
M. Ghirotto, P. H. Martins & R. Fonseca-Ferreira coll. (CAD 1202); Belo Horizonte,
Bairro São Bento, female, 17.II.2005, A. C. dos Santos coll. (UFMG 1774), Estação
Biológica da Universidade Federal de Minas Gerais, 19º52'26''S, 43º58'22''W, male,
05.X.2008, A. C. dos Santos coll. (UFMG 6087), male, 19º52'38''S, 43º58'16''W,
19.XI.2011, B. T. Faleiro et. Al coll. (UFMG 10082); Nova Lima, Condomínio
Alphaville, 20º4'0.16''S, 43º51'15''W, male, 11.XI.2009 (UFMG 6076), female (UFMG
6079), RPPN Mata Samuel de Paula, 20º00'S, 43º52'W, male, 14.X.2006, J. P. P. Pena-
Barbosa coll. (UFMG 2637).

Parvicarina Galleti-Lima, Hamilton, Borges and Guadanucci, new genus

Parvicarina felipeleitei (Bertani & Leal, 2016) new comb.
Pterinopelma sazimai Bertani, Nagahama & Fukushima, 2011: 8, figs. 7-16 (in part:
only the male, not examined). Pterinopelma felipeleitei Bertani & Leal, 2016: 4, figs. 8-
18, 20-22.
Diagnosis: See diagnosis for the genus.
Description: See Bertani et al. (2011) and Bertani & Leal (2016).
Additional material examined: BRAZIL, Minas Gerais: Diamantina, male, female,
05.VII.2011 (CAD 411); Gouveia, Fazenda Requeijão, 18º22'42.98S, 43º51'50.47''W,
female, VII.2019, P. H. Martins coll (UFMG 17211); Morro do Pilar, 19º13'14.82''S,
43º29'32.58''W, male, 21.IV.2012, E. A. Araújo coll. (UFMG 17203), female,
21.IV.2012, E . A. Araújo coll. (UFMG 17205);

Lasiocyano Galleti-Lima, Hamilton, Borges and Guadanucci, new genus.

Lasiocyano sazimai (Bertani, Nagahama & Fukushima, 2011) new comb.
Pterinopelma sazimai Bertani, Nagahama & Fukushima, 2011: 8, figs. 13-14, 16
(misidentified, belongs to P. felipeleitei, see Bertani & Leal, 2016; not examined).
Pterinopelma sazimai Bertani & Leal, 2016: 2, figs. 1-7, 19.
Diagnosis: See diagnosis for the genus.
Description: See Bertani et al. (2011) and Bertani & Leal (2016).
Additional material examined: BRAZIL, Bahia: Mucugê, road BA 245, 13º0'40.41''S,
41º24'35.73''W, female, 26.II.2017, P. H. Martins coll. (UFMG 24036); Minas Gerais:
Diamantina, 18°14'38.7"S, 43°35'52.9"W, male, 22.II.2004, G. A. Coelho coll. (UFMG
1769), Vila Operária, female, 30.IV.2009, Silvana coll. (CAD 003), male, XI. 2008
(CAD 004), male (CAD 084), female, Equipe de Zoonozes Prefeitura de Diamantina
coll. (CAD 087), female, Equipe de Zoonozes Prefeitura de Diamantina coll. (CAD
093), male, III. 2009 (CAD 095), female, XI.2008, Equipe de Zoonozes Prefeitura de

4
Diamantina coll. (CAD 096), female, Equipe de Zoonozes Prefeitura de Diamantina
coll. (CAD 098), male, XI. 2011 (CAD 442), male (CAD 466), male (CAD 479),
female (CAD 588), female, J. P. L. Guadanucci coll. (CAD 609), female (CAD 623),
Rua do Areião, Bairro Rio Grande, Equipe de Zoonozes Prefeitura de Diamantina coll.
(CAD 097), female, 24.X.2018, A. Galleti-Lima, V. M. Ghirotto, R. Fonseca-Ferreira
coll. (CAD 1269), 15.X.2018, A. Galleti-Lima, V. M. Ghirotto, R. Fonseca-Ferreira
coll. (CAD 1270), female (CAD 1272), female (CAD 1273); Lapinha da Serra, Pousada
Duas Pontes (Travessão trail), male, 17-21.X.2018, A. Galleti-Lima, P. H. Martins, V.
M. Ghirotto, R. Fonseca-Ferreira coll. (CAD 1260), female, 17-21.X.2018, A. Galleti-
Lima, P. H. Martins, V. M. Ghirotto, R. Fonseca-Ferreira coll. (CAD 1261), juvenile,
17-21.X.2018, A. Galleti-Lima, P. H. Martins, V. M. Ghirotto, R. Fonseca-Ferreira coll.
(CAD 1265), male, 17-21.X.2018, A. Galleti-Lima, P. H. Martins, V. M. Ghirotto, R.
Fonseca-Ferreira coll. (CAD 1267), road to Lapinha da Serra, juvenile, 17-21.X.2018,
A. Galleti-Lima, P. H. Martins, V. M. Ghirotto, R. Fonseca-Ferreira coll. (CAD 1262),
Parque Nacional da Serra do Cipó, juvenile, 17-21.X.2018, A. Galleti-Lima, P. H.
Martins, V. M. Ghirotto, R. Fonseca-Ferreira coll. (CAD 1266); São Gonçalo do Rio
das Pedras, female, 06.III.2013 (CAD 1271); Gouveia, Fazenda Fagundes,
18º25'35.67''S, 43º45'4.56''W female, XII.2018, P. H. Martins coll. (UFMG 17204),
male, 18.I.2011, P. H. Martins coll. (UFMG 17208), male, 15.XII.2015, P. H. Martins
coll. (UFMG 20074), male, 17.I.2011, P. H. Martins coll. (UFMG 17207), female,
25.XII.2015, P. H. Martins coll. (UFMG 20076), female, 18.I.2013, P. H. Martins coll.
(UFMG 17206), female, 25.XII.2015, P. H. Martins coll. (UFMG 20075), Fazenda
Requeijão, 18º23'48.54S, 43º51'9.06''W, female, 15-20.II.2012, P. H. Martins coll.
(UFMG 17209); Jaboticatuba, Parque Nacional da Serra do Cipó, 19°31'19.8"S,
43°44'43.6"W, male, 02.XI.2009, W. A. Fernandes coll. (UFMG 6249); Nova Lima,
Retiro do Chalé, 20°11'52.2"S, 44°00'09.0"W, male, 08.IX.2003, M. Bonjardim coll.
(UFMG 1304); Santana do Riacho, Fazenda Paláco, 19º16'39.72''S, 43º32'56.58''W,
male, 11.II.2017, F. Leal coll. (UFMG 23464), male, female, 17.VII.2012, P. H.
Martins coll. (UFMG 17202), female, VI.2011, E. A. Araújo coll. (UFMG 17212).

Tekoapora Galleti-Lima, Hamilton, Borges and Guadanucci, new genus

Tekoapora wacketi (Mello-Leitão, 1923) new comb.
Pterinopelma wacketi Mello-Leitao, 1923: 185 (holotype male from "Raiz da Serra",
São Paulo, Brazil, MZSP 147, 1900, M. Wacket col., not examined). Petrunkevitch,
1939: 261. Bonnet, 1958: 3828. Bücher, Costa & Lucas, 1971: 129. Pérez-Miles, 1992:
30. Eurypelma wacketi; Roewer, 1942: 242. Rhechostica wacketi; Raven, 1985: 158.
Aphonopelma wacketi; ICZN ruling, 1991: 166-167. Platnick, 1993: 100-101. 1997:
152. Pamphobeteus platyomma; Bücher, 1949: 117-135, figs. 1-3, T. 1-3
(misidentification). Pamphobeteus insularis Mello-Leitão, 1923: 241-242 (holotype
female from Ilha Grande, Rio de Janeiro, Brazil, should be in MNRJ, not located).
Petrunkevitch, 1939: 243. Bücher, 1949: 126 (syn. with Pamphobeteus platyomma).
Bonnet, 1958: 3314. Platnick, 1993:111. Syn. n. Pamphobeteus masculus Piza, 1939: 6
(holotype male from Ilha dos Alcatrazes, São Paulo, Brazil, MZLQ A0041, 1937,
Lepage col., not examined). Bücher, 1949: 126 (syn. with Pamphobeteus platyomma).
Brignoli, 1983: 139. Platnick, 1989: 126. Syn. n. Pamphobeteus litoralis Piza, 1976: 3
(holotype male from Registro, São Paulo, Brazil, ESALQ A-OI04, III11974, E. S.
Lopes col., not examined). Brignoli, 1983: 139. Lucas et aI., 1993: 245 (syn. with
Vitalius platyomma). Syn. n. Vitalius platyomma; Lucas et al., 1993: 241-245, figs. 1-3

5
(comb. n.). Platnick, 1997: 171. Vitalius wacketi Bertani, 2001: 294, figs. 16, 83-86,
179-180. V. wacketi Peters, 2003: 312, figs. 1250-1252, 1258-1260. V. wacketi Schmidt,
2003: 195, figs. 555-557.
Diagnosis: See diagnosis for the genus.
Description: See Bertani (2001).
Additional material examined: BRAZIL, Paraná: Matinhos, Parque Nacional Saint
Hilaire/Lange, female, 24.XI.2018, A. Galleti-Lima, E. F. Trova, R. Fonseca-Ferreira
coll. (CAD 1215); São Paulo: São Sebastião, male, 28.III.2005, G. Medlong coll. (MCN
38807), male (MCN 17741).

Pterinopelma Pocock, 1901

Pterinopelma Pocock 1901: 551 (type species by monotypy Eurypelma vitiosa
Keyserling 1891, holotype male, Brazil, Taquara, in BMNH, not examined); Mello-
Leitão 1923: 183, 1943: 153; Petrunkevitch 1928: 80; 1939a: 261; 1939b: 567; Bonnet
1958: 3827; Gerschman & Schiapelli 1978: 86; Valerio 1980: 276. Bertani et al. (2011):
5, figs. 1-16 (removed from the synonymy of Eupalaestrus Pocock, 1901).
Redescription: Carapace longer than wide. Cephalic region slightly raised (P. vitiosum)
to clearly raised (P. longisternalis and P. roseus). Cephalic and thoracic striae shallow,
convex. Fovea short and straight, shallow (P. vitiosum) or deep (P. longisternalis and P.
roseus). Chelicerae without rastellum, basal segments with 9 to 12 teeth. Eye tubercle
distinct, wider than long. Short clypeus or absent (female of P. vitiosum). Anterior eye
row procurved, posterior recurved. Labium wider than long, with numerous (100 to
200) cuspules. Maxilla with numerous cuspules (100 to 200). Sternum as long as wide
(P. vitiosum and some specimens of P. roseus) or longer than wide (P. longisternalis).
Sigilla not visible (P. vitiosum) ou anterior sigilla on the sternum/labium edge and other
sigilla submarginal, the second pair smaller than the third that is smaller than the fourth
(P. longisternalis and P. roseus). Leg formula: IV I II III. Clavate trichobothria on distal
2/3 of tarsi I–IV. Absense of stridulatory apparatus. Femur IV with retrolateral scopula.
Leg tarsi without spines, claw tufts present. Tarsi I–III fully scopulated, IV divided by
row of setae (P. vitiosum) or tarsi I–III fully scopulated (P. longisternalis and P.
roseus). Metatarsi I-II scopulated, metatarsus III 1/3 or 1/2 scopulated. Metatarsus IV
apically scopulated. Type I and III urticating hair present in males. Only type I
urticating hair present in females. Type I urticating setae in females with region "A"
longer or equal to region "B". Male palpal bulb pyriform, embolus slightly flattened
distally. Embolus long. Prolateral keels present, the Prolateral Superior keel forming the
embolus apical edge. Acessory keel absent. Retrolateral keel present, not pronounced
(P. vitiosum) or pronounced, sharp (P. longisternalis and P. roseus). Apical keel small
(P. vitiosum) or medially developed (P. longisternalis and P. roseus). Subapical keel
present, but inconspicuous (P. vitiosum), well developed, bordered by small deticles (P.
longisternalis) or small (P. roseus). Male spur with converging branches originating
from a common base, tapering distally, the prolateral branch straight (P. vitiosum and P.
longisternalis) or male spur branches very small, originating from a common base (P.
roseus). Metatarsus I slightly curved, when folded touches retrolateral tibial apophysis
branch laterally (P. vitiosum), straight, when flexed touches the retrolateral branch
laterally (P. longisternalis) or straight, when flexed passes by the retrolateral branch
Iaterally without touching it (P. roseus). Small spermathecae with receptacles separated

6
by a small central depression and spermathecal stalk with a slight constriction at the
base of the spermathecal bulb. Carapace covered by short slender setae, bordered by
short setae pointing out. Legs ventrally covered by many long setae. Coxae and sternum
covered by long (P. vitiosum) or short (P. longisternalis and P. roseus) setae. Color
pattern ranging from homogeneous brown becoming darker on the cephalic region,
marginated by light brown (P. vitiosum and P. roseus) or dark, margin lighter (P.
longisternalis). Coxae, labium, sternum, maxila and legs dark brown (P. vitiosum and P.
roseus) or dark (P. longisternalis). Legs dark brown with light brown long setae (P.
vitiosum), dark (P. longisternalis) or with trochanters dorsally light brown, femora
black, patellae, tibiae and metatarsi of the legs I and II laterally pinkish (P. roseus). Leg
rings and stripes inconspicuous (P. vitiosum), leg rings on the apex of trochanter and
femora and stripes easly distinct (P. longisternalis) or leg rings very distinct on the apex
of the femora, patellae, and tibiae, with longitudinal leg stripes distinct, pinkish (P.
roseus). Abdomen dorsally black.

Pterinopelma vitiosum (Keyserling, 1891)

Eurypelma vitiosum Keyserling 1891: 21, pl. 1, f.5 (holotype male from Brazil,
Taquara, in BMNH, not examined); Simon 1903: 937; Petrunkevitch 1911: 65, 776;
Roewer 1942: 242, 1954: 1508. Eurypelma mollicomum: Simon 1892: 167 nec
Ausserer, 1875. Pterinopelma vitiosum: Pocock 1901: 551, 1903: 108; Mello-Leitão
1923: 190, 1943: 153; Petrunkevitch 1928: 80, 1939: 261; Bonnet 1955: 1831;
Capocasale 1980: 5. Eupalaestrus vitiosum: Pérez-Miles 1992: 34; Platnick, 2010. First
considered as nomen dubium by Pérez-Miles 1992: 34. Pterinopelma vitiosum; Bertani,
Nagahama & Fukushima, 2011: 5, figs. 1-16.
Diagnosis: Males of P. vitiosum differ from congeneres by the presence of a narrow and
inconspicuous subapical keel. Females of P. vitiosum differ by the shape of the rounded
sternum when compared to the sternum of P. longisternalis (longer than wide) and P.
roseus (not rounded).
Description: See Bertani et al. (2011)
Additional material examined: BRAZIL, Rio Grande do Sul: Caxias do Sul,
Universidade de Caxias do Sul, female (CAD), Cânions Palanquinhos, juvenile (CAD);
female (MCN 22102); Encanto, 29º14’02.8’’S, 51º52’10.5’’W, male, 17.IV.1992, L.
Dacroce coll. (MCN 22145).

Pterinopelma longisternalis (Bertani, 2001) new comb.

Vitalius longisternalis Bertani, 2001: 297, figs. 21, 87-90, 181 (Holotype Male, IBSP
6772, Candi, Paraná, Brazil, IV/1995, Faunal Rescue Team of Hydroelectric Power
Station of Segredo/derivação do Rio Jordão, col., not examined. Paratype. Female, IBSP
7017, same data of holotype, not examined). Vitalius longisternalis; Peters, 2003: 307,
figs. 1215, 1218-1219. V. longisternalis; Schmidt, 2003: 193, figs. 536-538. V.
longisternalis; Peters, 2005: 120, figs. 382-387. V. longisternalis; Ferretti et al., 2015: 1,
figs. 1a-b, 5.
Diagnosis: Males can be distinguished from Pterinopelma vitiosum by the presence of a
long subapical keel and from Pterinopelma roseus by a long male spur (Figures 9D-F).

7
Females can be distinguished by the sternum longer than wide (except for some
population of Pterinopelma roseus, that occurs strictly in Atlantic Forest) (See Bertani,
2001, Fig. 21). From these individuals, they differ by the collor pattern, with carapace
dark, margin lighter, sternum, coxae and abdomen ventrally dark and leg rings on the
apex of the femora (Figure 8C).
Description: See Bertani (2001).
Additional material examined: BRAZIL, Paraná: Altônia, Parque Nacional de Ilha
Grande, female, (IBSP 210343); Andirá, female, R. L. Dirickson coll. (IBSP 106661);
Cruzeiro do Iguaçú, Foz do Chopim (Usina Hidrelétrica Salto Caxias), juvenile (IBSP
108132), female (IBSP 108133), female (IBSP 108134), female (IBSP 108139), female
(IBSP 108141), female (IBSP 108143), female (IBSP 108144), female (IBSP 108146),
female (IBSP 108148), female (IBSP 108149), female (IBSP 108150), female (IBSP
108160), female (IBSP 108162), male (IBSP 108364), female, 15.X.1997, R. S.
Bennils, S. A. A. Morato & G. G. Montingelli coll. (MZUSP 74375); Cruzeiro do
Oeste, Foz do Chopin, female, 23.III.1993, Morato & Moma-Leite coll. (MCN 23409);
Fernandes Pinheiro, Floresta Nacional de Iratí, female, 04.XII.2020, W. Moeller, H.
Gonzalez-Filho & A. Galleti-Lima coll. (CAD 1212), juvenile (CAD 1213), juvenile
(CAD 1214); Foz do Iguaçú, male (MZUSP 74055); Guarapuava, male, W. Moeller
coll. (CAD 1240), male, W. Moeller coll. (CAD 1240), male, W. Moeller coll. (CAD
1242), male, 2019, W. Moeller coll. (CAD 1247); Laranjeiras do Sul, male, W. Moeller
coll. (CAD 1239); Pinhão, Usina Hidrelétrica de Santa Clara, female (IBSP 151210);
Pitanga, Fazenda Vale Verde, male, 2019, N. Porfírio coll. (CAD 1211); Três Barras do
Paraná, Flor da Serra (Usina Hidrelétrica Salto Caxias), female (IBSP 108152), female
(IBSP 108153), female (IBSP 108155), female (IBSP 108156), female (IBSP 108159),
female (IBSP 108161); Santa Catarina: São Domingos, male, 05-13.IV.2001, M.
Krammers (MCTP 11344); Urubici, Parque Nacional de São Joaquim, 11-18.I.2019, J.
P. L. Guadanucci, R. P. Indicatti, B. Gambaré, R. Fonseca-Ferreira & A. Galleti-Lima
coll. (CAD).

Pterinopelma roseus (Mello-Leitão, 1923) new comb.

Pamphobeteus roseus Mello-Leitão, 1923: 232 (holotype female from Itaquy (=Itaqui),
Rio Grande do Sul, Brazil, MZSP 143, 1914, E. Garbe col., not examined).
Petrunkevitch, 1939: 244. Bücherl, 1947a: 257 (in part). Bonnet, 1958: 3315. Schmidt,
1986: 59, fig. 97 (in part). Platnick 1989: 107. Platnick, 1993: III. Vitalius roseus Lucas,
Silva Junior & Bertani, 1993: 241-245 (comb. n.). Platnick, 1997: 171. V. roseus;
Bertani, 2001: 305, figs. 112-116, 185. V. roseus; Peters, 2003: 309, figs. 1228-1231,
1233-1234. V. roseus; Schmidt, 2003: 195, figs. 548-550. V. roseus; Ferretti et al.,
2015a: 241, figs. 1A-D, 2A-B, 5A-B.
Diagnosis: Males can be distinguished from congeneres by the presence of a small,
male spur (Figure 9I). Females can be distinguished from Pterinopelma vitiosum by the
shape of sternum, not rounded or longer than wider (See Bertani et al., 2011, Fig. 5) and
leg rings and stripes very distinct (Fig. 8E). Individuals with sternum longer than wide
differ from Pterinopelma longisternalis by the color pattern on sternum, coxae and
abdomen being black ventrally and patellae and metatarsi dorsolaterally slightly pinkish
(Figure 8E).
Description: See Bertani (2001).

8
Additional material examined: BRAZIL, Rio Grande do Sul, Bossoroca, female,
14.VII.1973, A. Lise coll. (MCN 1741); Canela, female (MCTP 9939); Carazinho,
male, 23.V.1974, R. Bohrer (MCN 4011); Cerro Largo, Fundão do Ijui, male, L. M.
Borges coll. (CAD LMB 075), Cerro Largo, Fundão do Ijui, juvenile, L. M. Borges coll.
(CAD LMB 188); Cruz Alta, Fazenda Umbu, female, 10-22.VII.1976, F. Silva coll.
(MCN 4357); Derrubadas, close to Parque Estadual do Rio Turvo, female, L. M. Borges
coll. (CAD LMB 149), male, M. Rocha coll. (CAD LMB 150); Esmeralda, female,
05.IX.2005, Bourscheld coll. (MCN 42460); Iraí, female, 19.XI.1975, A. Lise coll.
(MCN 3045); Itaara, male (MCTP 39401); Itauba, Arroio do Tigre, female, 11.IV.1978,
A. Lise coll. (MCN 17731); Jaguari, Chacara Dri, female, L. M. Borges coll. (CAD
LMB 018), female, L. M. Borges coll. (CAD LMB 045), female, L. M. Borges coll.
(CAD LMB 018), female, L. M. Borges coll. (CAD LMB 110), juvenile, L. M. Borges
coll. (CAD LMB 111), female, L. M. Borges coll. (CAD LMB 145), female, L. M.
Borges coll. (CAD LMB 146), male, L. M. Borges coll. (CAD LMB 147), Instituto
Federal Farroupilha, female, L. M. Borges coll. (CAD LMB 043); Nova Palma, Rincão
do Padilha, male, C. Mario de Rosa coll. (CAD LMB 138); Panambi, Instituto Federal
Farroupilha, male, L. M. Borges coll. (CAD LMB 134), male, L. M. Borges coll. (CAD
LMB 136); Paraíso do Sul, female, R. Sá coll. (CAD LMB 148); Rosário do Sul,
Caverazinho (Serra do Caverá), female, L. M. Borges coll. (CAD LMB 009), female, L.
M. Borges coll. (CAD LMB 034), male, L. M. Borges coll. (CAD LMB 073), juvenile,
L. M. Borges coll. (CAD LMB 104), female, L. M. Borges coll. (CAD LMB 133);
Santa Maria, male, 18.IV.1980, D. Link coll. (MCN 9549), male, 18.IV.1980, D. Link
coll. (MCN 9550), Universidade Federal de Santa Maria, female, L. M. Borges coll.
(CAD LMB 046), female, L. M. Borges coll. (CAD LMB 066), juvenile, L. M. Borges
coll. (CAD LMB 105), female, L. M. Borges coll. (CAD LMB 112), juvenile, L. M.
Borges coll. (CAD LMB 128), female, L. M. Borges coll. (CAD LMB 143), male, L.
M. Borges coll. (CAD LMB 196), female, L. M. Borges coll. (CAD LMB 203), male,
L. M. Borges coll. (CAD LMB 396), male, L. M. Borges coll. (CAD LMB 399), male,
L. M. Borges coll. (CAD LMB 403), Monumento ao Ferroviário (Itararé), female, L. M.
Borges coll. (CAD LMB 049), female, L. M. Borges coll. (CAD LMB 050), female, L.
M. Borges coll. (CAD LMB 057), male, female, L. M. Borges coll. (CAD LMB 201),
juvenile, L. M. Borges coll. (CAD LMB 444), female, L. M. Borges coll. (CAD LMB
445), female, L. M. Borges coll. (CAD LMB 445), female, L. M. Borges coll. (CAD
LMB 446), juvenile, L. M. Borges coll. (CAD LMB 447), Morro do Elefante, male, L.
M. Borges coll. (CAD LMB 330), Highway BR287, female, L. M. Borges coll. (CAD
LMB 067), male, L. M. Borges coll. (CAD LMB 398); Santiago, male (MCTP 11202);
São Borja, Garruchos, 07.XI.1979, H. Bischoff coll. (MCN 9998); São Francisco de
Paula, female (MCTP 4850); São Martinho da Serra, 1 kilometer of municipality,
juvenile, C. Mario Rosa coll. (CAD LMB 432), juvenile, C. Mario Rosa coll. (CAD
LMB 433), juvenile, C. Mario Rosa coll. (CAD LMB 434); São Vicente do Sul,
Instituto Federal Farroupilha, female, L. M. Borges coll. (CAD LMB 209); Tenente
Portela, female, 10.IX.1976, S. Scherer coll. (MCN 4524), Parque Estadual do Rio
Turvo, female, 06.II.1980, T. de Lima coll. (MCN 8987), male (MCTP 10072); Três
Passos, female, A. D. Abegg coll. (CAD LMB 194); Tuparendí, female, 02.VII.1991, E.
Lückemeyer coll. (MCN 21232), female, 30.IV.1991, E. Lückemeyer coll. (MCN
21031), male, 07.V.1989, E. Lückemeyer coll. (MCN 21075), male, 20.V.1991, E.
Lückemeyer coll. (MCN 21197), male, 20.V.1991, E. Lückemeyer coll. (MCN 21200),
male, 20.V.1991, E. Lückemeyer coll. (MCN 21199), male, 20.V.1991, E. Lückemeyer
coll. (MCN 21198), male, 20.V.1991, E. Lückemeyer coll. (MCN 21209), female,
20.V.1991, E. Lückemeyer coll. (MCN 21210), male, 07.V.1989, E. Lückemeyer coll.

9
(MCN 21074), male, 30.IV.1991, E. Lückemeyer coll. (MCN 21027), male,
30.IV.1991, E. Lückemeyer coll. (MCN 21025), male, 30.IV.1991, E. Lückemeyer coll.
(MCN 21026), male, 30.IV.1991, E. Lückemeyer coll. (MCN 21028), male,
30.IV.1991, E. Lückemeyer coll. (MCN 21024), female, 30.IV.1991, E. Lückemeyer
coll. (MCN 21029), female, 30.IV.1991, E. Lückemeyer coll. (MCN 21030), male,
07.V.1989, E. Lückemeyer coll. (MCN 21076);

Vitalius Lucas, Silva & Bertani, 1993

Pamphobeteus Pocock, 1901:545 (in part). Mello-Leitão, 1923:225. Vitalius Lucas,
Silva Junior & Bertani, 1993:241, figs. 1-3; type species by original designation Vitalius
sorocabae Mello-Leitão, 1923. Pérez-Miles et al., 1996:59, figs. 49-50. Platnick, 1997:
170. Pterinopelma Pocock, 1901:551 (in part). Mello-Leitão, 1923: 185-189.
Rhechostica Simon, 1892:162 (in part). Raven, 1985:158 (syn. Pterinopelma).
Aphonopelma Pocock, 1901:553 (in part). Platnick, 1993:100.
Redescription: Cephalothorax longer than wide, cephalic region slightly raised,
convex. Cephalic and thoracic striae distinct (most species) or hardly distinct (V.
sorocabae). Fovea short, deep, straight, slightly recurved or procurved. Chelicerae
without rastellum, basal segments with 10 to 15 teeth. Eye tubercle distinct, wider than
long. Clypeus absent or short. Labium slightly wider than long, with numerous cuspules
(100 to 200). Maxila subretangular, anterior lobe distinctly produced into a conical
process, inner angle bearing numerous cuspules (100 to 200). Sternum longer than wide
(female of Vitalius sp. nov.) or as long as wide (other species). Anterior sigilla on the
sternum/labium edge. Other sigilla submarginal, the second pair smaller than the third
that is smaller than the fourth or only the third and fourth pair visible (Vitalius sp. nov).
Leg formula: IV I II III. Absense of stridulatory apparatus. Leg tarsi without spines,
claw tufts present. Tarsi I-IV and metatarsi I-II scopulated, metatarsus III scopulated
along half its length, metatarsus IV apically scopulated. Femur IV with retrolateral
scopula. Type I and III urticating hair present in males. Only type I urticating hair
present in females. Type I urticating setae in females with region "A" longer or equal to
region "B". Male spur with converging branches originating from a common base,
tapering distally, the prolateral branch thickened (most species) or straight branches
originating from a common base, the retrolateral constricted in the middle (V. lucasae)
or male spur vestigial (some morphotypes of V. vellutinus). Metatarsus I curved (V.
lucasae) or straight (other species), when flexed touches the retrolateral branch laterally.
Male palpal bulb pyrifonn, embolus slightly flattened distally. Embolus long (Vitalius
sp. nov.) or short (other species). Prolateral keels present, the Prolateral superior
forming the embolus edge distally, pronounced in V. dubius, V. buecherli and Vitalius
sp. nov. Accessory keel absent. Retrolateral keel present, pronounced, sharp. Apical
keel present, medially developed. Subapical keel well-developed, triangular, bordered
by small denticles, or vestigial (V. vellutinus). Male palpal bulb with accentuated (V.
lucasae) or slightly (other species) ventral median depression. Spermathecae short,
separated by a heavily sclerotized short area or small spermatheca homogeneously
sclerotized (Vitalius sp. nov.) Spermathecal stalk narrower than spermathecal bulb,
with elevated central region (Vitalius sp. nov.). Carapace covered by short slender setae;
bordered by some longer setae ponting to the carapace center (V. sorocabae female) or
bordered by short setae pointing out (V. sorocabae male and other species). Tibiae IV
slightly thickened in male and female (V. lucasae) or tibiae IV not thickened (other
species). Legs dorsally and ventrally covered by few long setae (females of V. vellutinus

10
and V. paranaensis) or short setae (Vitalius sp. nov.) or covered by many long setae,
mainly on the ventral face (other species). Coxae and sternum covered by long setae (V.
buecherli female) or short (other species).

Vitalius australis Galleti-Lima, Hamilton, Borges and Guadanucci, new species

Additional material examined: BRAZIL, Paraná: Ponta Grossa, male and female,
31.I.1975, I. Wagner coll. (IBSP 4195).
Description: Male holotype (CAD 1244). Total length: 34.2. Carapace 14.5 long, 13.0
wide. Chelicerae 3.6. Eye tubercle 1.3 long, 2.2 wide. Labium 2.0 long, 2.7 wide.
Sternum 6.5 long, 5.5 wide. Fovea short, deep, straight. Cheliceral basal segments with
12 teeth. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 12.8, 6.9, 10.4, 9.4, 6.6,
46.1. II: 11.9, 5.9, 7.4, 7.2, 6.4, 38.8. III: 10.3, 5.0, 7.6, 9.0, 5.7, 37.6. IV: 13.5, 6.0,
11.0, 10.2, 7.2, 47.9. Palp: 8.3, 4.3, 6.7, -, 2.5, 21.8. Spines: tarsi lacking spines. Palpal
femur p0, 0, 1, patela 0, tibia p1, 2, 2ap; legs I femur 0, patella 0, tibia v1, 2, 0ap, r0, 0,
1ap, metatarsus v0, 0, 1ap; II femur 0, patela 0, tibia v2, 2, 3ap, p0, 2, 1, r0, 0, 1,
metatarsus v1, 0, 3ap, p0, 1, 0; III femur 0, patela 0; tibia v1, 4, 4ap, p1, 3, 10ap, r0, 2,
2, d0, 0, 1, metatarsus v0, 1, 3ap, p3, 2, 1, r0, 2, 1, d0, 0, 1; IV femur 0, patela 0, tibia
v0, 4, 2ap, p1, 2, 1, r0, 2, 1, metatarsus v1, 9, 4ap, p1, 4, 1, r0, 5, 1, d0, 0, 1. Male spur
branches originating from a common base, retrolateral branch tapering distally and
longer than prolateral, short with rounded apex. Metatarsus I straight, when flexed
touches the retrolateral branch laterally. Male palpal bulb pyriform, long embolus,
slightly flattened laterally at the distal region. Prolateral keels present, the prolateral
superior forming the embolus edge distally. Rretrolateral present, pronounced, sharp.
Apical keel present, medially developed. Subapical well-developed, triangular. Types I
and III urticating setae present, type III restricted to a small area of the abdomen. Legs,
coxae and sternum covered by short setae ventrally. Carapace dark brown, bordered by
light brown setae. Legs dark, except trochanters, that are dorsally light brown. Sternum,
coxae and abdomen ventrally dark brown. Leg rings on the apex of the femora, patellae
and tibiae very distinct. Longitudinal leg stripes low distinct except for the tibiae stripes.
Legs, coxae and sternum covered by short setae ventrally.
Female paratype (CAD 1210). Total length: 54.0. Carapace 22.6 long, 18.3 wide.
Chelicerae 6.7. Eye tubercle 2.3 long, 3.3 wide. Labium 3.2 long, 4.2 wide. Sternum
10.5 long, 7.7 wide. Fovea short, deep, straight. Cheliceral basal segments with 12 teeth.
Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 14.6, 9.4, 10.3, 8.4, 6.5, 49.2. II:
13.1, 8.0, 7.3, 7.0, 5.6, 41,0. III: 11.5, 7.5, 7.6, 8.2, 4.8, 39,6. IV: 14.0, 7.3, 10.1, 12.3,
4.6, 48.3. Palp: 10.5, 6.6, 6.8, -, 7.4, 31.3. Spines: tarsi lacking spines. Palpal femur 0,
patela 0, tibia v0,0, 1ap, p0, 2, 1, r0, 0, 2; legs I femur 0, patella 0, tibia v0, 0, 2ap, p0,
1, 1ap, metatarsus v0, 0, 1ap; II femur 0, patela 0, tibia v1, 1, 2ap, p0, 1, 1, metatarsus
v0, 0, 1ap; III femur 0, patela 0, tibia v0, 0, 1ap, p0, 2, 0, r1, 1, 1, metatarsus v1, 1, 1ap,
p1, 1, 1, r1, 1, 1; IV femur 0, patela 0, tibia v0, 0, 1ap, p0, 2, 0, r1, 1, 1, metatarsus v3,
6, 1ap, p1, 1, 1, r0, 1, 0, d0, 0, 1. Small spermatheca homogeneously sclerotized.
Spermathecal stalk with a strong constriction at the base of the spermathecal bulb Types
I urticating setae present, type III not present. Legs, coxae and sternum covered by short
setae ventrally. Carapace brown, bordered by light brown setae. Legs light brown,
except femurs and trochanters, which are black dorsally. Sternum, thigh and abdomen
ventrally black.

11
Other paratypes - Male paratype (CAD 1243). Total length: 36.6. Carapace 16.2 long,
14.4 wide. Chelicerae 4.6. Eye tubercle 1.6 long, 2.3 wide. Labium 2.1 long, 3.0 wide.
Sternum 7.6 long, 6.3 wide. Fovea short, deep, straight. Cheliceral basal segments with
12 teeth. Legs (femur, patella, tibia, metatarsus, tarsus, total): I: 14.7, 7.6, 10.8, 9.8, 7.0,
49.9. II: 13.6, 7.0, 9.9, 9.0, 6.8, 46.3. III: 10.7, 5.8, 9.0, 10.3, 6.4, 42.2. IV: 14.4, 6.5,
9.1, 14.9, 6.7, 51.6. Palp: 8.6, 4.8, 6.9, -, 3.2, 23.5. Spines: tarsi lacking spines. Palpal
femur 0, patela p1, tibia v0, 1, 0ap, p0, 2, 1, d0, 0, 2ap; legs I femur d0, 0, 1, patella 0,
tibia v0, 2, 0ap, r1, 0, 0, metatarsus v0, 0, 2ap; II femur 0, patela 0, tibia v1, 2, 2ap, p0,
0, 1ap, d0, 1, 0, metatarsus v0, 2, 2ap, p0, 1, 0ap; III femur 0, patela 0, tibia v0, 4, 2ap,
p1, 1, 1ap, r1, 1, 1, metatarsus v1, 3, 3ap, p1, 2, 1, r0, 1, 1; IV femur 0, patella 0, tibia
v0, 4, 2ap, p0, 3, 0ap, r0, 2, 1ap, metatarsus v3,8, 2ap, p0, 1, 1ap, r0, 1, 1ap. Female
paratype (CAD LMB). Total length: 48.0. Carapace 19.3 long, 16.4 wide. Chelicerae
7.3. Eye tubercle 2.0 long, 2.6 wide. Labium 2.4 long, 3.9 wide. Sternum 10.4 long, 7.2
wide. Fovea short, deep, straight. Cheliceral basal segments with 12 teeth. Legs (femur,
patella, tibia, metatarsus, tarsus, total): I: 14.1, 8.5, 9.5, 8.1, 5.4, 45.6. II: 11.8, 8.0, 8.4,
7.3, 5.0, 40.5. III: 10.0, 7.3, 7.1, 8.5, 4.5, 37.4. IV: 13.3, 7.4, 9.9, 12.4, 5.4, 48.4. Palp:
10.3, 6.5, 7.2, -, 6.6, 30.6. Spines: tarsi lacking spines. Palpal femur p0, 0, 1ap, patella
0, tibia v0,0, 2ap, p0, 2, 1; legs I femur p0, 0, 1, patella 0, tibia v0, 0, 2ap, p1, 1, 1,
metatarsus v1, 0, 1ap; II femur p0, 0, 1ap, patella 0, tibia v0, 1, 2ap, p1, 1, 1, metatarsus
v1, 0, 3ap, p1, 1, 0; III femur r0, 0, 1ap, patella r1, tibia v0, 1, 1ap, metatarsus p0, 5,
1ap, r1,1,1; IV femur d0, 0, 1ap, patella 0, tibia v0, 2, 1ap, p0, 1, 1ap, r1, 1, 1,
metatarsus v3,4,4ap, p1, 2, 1ap, r0, 1, 1, d0, 0, 1ap.

12