Tityus forcipula
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species group
References after Fet & Lowe (2000: 230). Armas et al. (2002): 166 [in part]; Lourenço & Leguin (2008): 3; Teruel & García (2008): 1; Kovařík et al. (2013): 8 [in part]; Lourenço & Ythier (2013): 3; Brito & Borges (2015): 7, 8, 9, 14 [in part], fig. 4G; Flórez (2015): 85; Lourenço (2016): 7 [in part]; Miranda et al. (2020): 197, fig. 27.
Included species.
Tityus crassicauda
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,
Tityus cuellari
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,
Tityus forcipula
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,
Tityus fuhrmanni
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, and
Tityus moralensis
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sp. nov.
Excluded species.
Tityus dasyurus fulvipes
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,
Tityus festae
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,
Tityus florezi
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,
Tityus jaimei
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,
Tityus macrochirus
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,
Tityus metuendus
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, and
Tityus pachyurus
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are here transferred to the
Tityus obscurus
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species group.
Tityus dasyurus dasyurus
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is excluded from the
Tityus forcipula
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species group, but not assigned to any species group, until additional morphological evidence is gathered.
Amended diagnosis. The
Tityus forcipula
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species group is more similar to the
Tityus bahiensis
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,
Tityus trivittatus
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, and
Tityus stigmurus
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species groups, and differs from
Tityus androcottoides
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,
Tityus bolivianus
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,
Tityus clathratus
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, and
Tityus obscurus
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species groups. The
Tityus forcipula
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,
Tityus bahiensis
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,
Tityus trivittatus
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, and
Tityus stigmurus
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species groups share ventral macrosetae of telotarsi I–IV distributed in two submedian rows (type II). Whereas, the
Tityus androcottoides
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,
Tityus bolivianus
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,
Tityus clathratus
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, and
Tityus obscurus
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species groups exhibit ventral setae of telotarsi I–IV irregularly distributed in a tuft (type I).
On the other hand, the
Tityus forcipula
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species group differs from the
Tityus bahiensis
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,
Tityus stigmurus
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and
Tityus trivittatus
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species groups in the following character state combination: i) pectines strongly sclerotized and brown colored ( Figures 2B, D
View FIGURE 2
, 3B, D
View FIGURE 3
, 9A–D
View FIGURE 9
); ii) basal middle lamellae of female pectines dilated and suboval shaped or ovoid ( Figures 2D
View FIGURE 2
, 3D
View FIGURE 3
, 9B, D
View FIGURE 9
); iii) female pectinal basal piece without glandular region; iv) ventral macrosetae of telotarsi I–IV strongly sclerotized and stout; v) metasomal segments (moderately or strongly) widening towards segment V ( Figures 12A–D
View FIGURE 12
, 13A–D
View FIGURE 13
) (except in
Tityus fuhrmanni
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that exhibit a conspicuous thinning towards segment V); and vi) subaculear tubercle small to medium and coarse with a small gap between the dorsal margin of the subaculear tubercle and the base of the aculeus ( Figure 15A–D
View FIGURE 15
). Whereas,
Tityus bahiensis
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,
Tityus stigmurus
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and
Tityus trivittatus
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species groups present i) pectines moderately sclerotized and white colored; ii) basal middle lamellae of female pectines not dilated; iii) female pectinal basal piece with a moderate to large glandular region; iv) ventral setae of telotarsi I–IV moderately sclerotized and fine; v) metasomal segments of similar wide or with an inconspicuous widening towards segment V, and vi) subaculear tubercle small and acute with a moderate gap between the dorsal margin of the subaculear tubercle and the base of the aculeus.
Taxonomic remarks. The
Tityus forcipula
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species group was previously merged with the
Tityus obscurus
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species group (previously referred as
T. asthenes
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, see Lourenço et al. (2019)). However, based on a unique combination of phenotypic characters here is revalidated and separated from the
Tityus obscurus
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species group. In order to set the boundaries of the
Tityus forcipula
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species group we examined all the species that belong to this group, but
Tityus crassicauda
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and
T. dasyurus dasyurus
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.
In our morphological survey, we found that
Tityus dasyurus fulvipes
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,
Tityus festae
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,
Tityus florezi
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,
Tityus jaimei
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,
Tityus macrochirus
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,
Tityus metuendus
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, and
Tityus pachyurus
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, exhibit ventral setae of telotarsi I–IV irregularly distributed in a tuft (type I). This character state together with the presence of pectines moderately sclerotized and white colored and basal middle lamellae of female pectines dilated and subcircular, exclude these taxa from the
Tityus forcipula
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species group and locate them into the
Tityus obscurus
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species group. Other characters such as a metasoma progressively widening towards segment V (club-like shaped metasoma) are shared by
Tityus jaimei
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,
T. metuendus
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, and
T. pachyurus
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, as well as, species belonging to the
Tityus forcipula
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species group. However, this character by itself is not enough to assign species to a given species group and must be used in combination with other characters with strong phylogenetic signal (e.g., Moreno-González et al. 2021).
Lourenço & Ythier (2013) described
Tityus crassicauda
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and mentioned that it is closely related to
T. forcipula
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. Their description has a detailed enough description that allows us to identify most of the character states of the
Tityus forcipula
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species group. For example, the authors described the presence of two ventrosubmedian rows in the telotarsi (type II), the metasoma widening towards segment V and the dilation and oval shape of the basal middle lamellae of female pectines. Also, they provided colorful habitus pictures of both sexes that clearly show the strongly sclerotized pectines and the absence of glandular regions in the pectinal basal piece of the female. The situation of
Tityus dasyurus
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is, however, diametrically opposite.
Pocock (1897) described
Tityus dasyurus
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based on a single female from “ Porto Rico ”. Nevertheless, some authors argued it corresponds to a mislabelling error and this female most probably comes from a locality in Central or South America ( Armas 2020). Santiago-Blay (1985) examined and redescribed the female holotype of
T. dasyurus dasyurus
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. In this description, we observed that the morphology of the subaculear tubercle and the basal middle lamellae shape are very different from other members of the
Tityus forcipula
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species group. The subaculear tubercle is conical, small and acute and presents a moderate gap between the dorsal margin of the subaculear tubercle and the base of the aculeus ( Santiago-Blay 1985: fig. 13), whereas the basal middle lamellae is dilated and subcircular shaped ( Santiago-Blay 1985: fig. 10). Both characters are very common among the
Tityus androcottoides
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and
Tityus obscurus
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species group members. However, the examination of additional morphological characters is needed to confirm the species group membership of this enigmatic species.
Finally,
Tityus fuhrmanni
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is particularly interesting. This species was assigned to an independent species group by Mello-Leitão (1945) given the particular morphology of its metasoma, which exhibits a conspicuous and progressive thinning towards segment V, but also presents a remarkable development of the DSM on the metasomal segment III. We also noted that the males of
T. fuhrmanni
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do not exhibit a bulkier chela compared to female chela, both male and female chelae ratios are somewhat similar. In this contribution, we temporarily followed the Lourenço (1984) designation of
T. fuhrmanni
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in the
T. forcipula
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species group. This species shares some character states with this species group such as i) telotarsi macrosetae stout and distributed in two ventrosubmedian rows (type II), ii) sclerotized pectines brown colored, iii) basal middle lamellae dilated and soboval/ovoid shaped in the female pectines, and iv) absence of glandular regions in the basal pectinal piece of the female pectine. However, additional studies and samples are required for testing the membership of this species into the
Tityus forcipula
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species group.