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Molecular phylogenetic tree for family Mithracidae sensu stricto based on 9 genera and 36 species in concatenated analysis of 3 mitochondrial genes (12S, 16S, COI), represented as maximum likelihood phylogram with maximum likelihood bootstrap values and Bayesian posterior probabilities. (* = 100% support, ¤ = 50% support). 

Molecular phylogenetic tree for family Mithracidae sensu stricto based on 9 genera and 36 species in concatenated analysis of 3 mitochondrial genes (12S, 16S, COI), represented as maximum likelihood phylogram with maximum likelihood bootstrap values and Bayesian posterior probabilities. (* = 100% support, ¤ = 50% support). 

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Mithracid crabs comprise a primarily subtidal reef- and rubble-dwelling group inhabiting both tropical and subtropical seas. Despite their relative ubiquity in many hard-substrate environments, there has been little consensus about their phylogenetic relationships or whether their group rank should be that of subfamily or family. We have used a com...

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... is also included within Mithracidae on the basis of phylogenetic analyses (>95/>95) ( Fig. 1). Morphological examination of specimens from the six species included in the genetic analysis also supports this placement on the basis of relative development of the ambulatory leg dactylar locking mechanism and the expanded orbital floor typical of mithracids. The generic grouping is very well supported (100/ 100) (Fig. ...
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... our global review of mithracid genera, based on a combination of morphological and molecular genetic data, present evidence suggests that Mithracidae is an amphi- American family with a single known outlying representative in the eastern Atlantic (Mithrax caboverdianus). In the recent synopsis by Ng et al. (2008), 17 genera and 108 species are placed within Mithracinae (our Mithracidae), but independent analyses of the superfamily Majoidea excludes five of those genera (Macrocoeloma, Micippa, Picroceroides, Stenocionops, and Tiarinia) and incorporate two others (Hemus and Pitho) (Figs 1, 2; Windsor, Ahyong and Felder, unpubl. data). Thus, the revised composition of Mithracidae used in our present work includes 13 genera and 36 species (Fig. 2). The present results support the naming or resurrection of four genera (three among species most recently assigned to Mithrax) and require removal of seven species from Mithrax, four from Microphrys, and one from Mithraculus, with placement of them into newly erected or alternative existing genera (see Systematic ...
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... Mithracidae s.s., Mithrax (sensu Ng et al. 2008) is a polyphyletic genus that can be reconfigured into five genera, three of which warrant new names and treatment under new diagnoses. Mithrax s.s., defined by the type species M. aculeatus (Herbst, 1790), groups most closely with M. hemphilli Rathbun, 1892 and M. caboverdianus Türkay, 1986 though with high support (92/100). The latter of these represents the sole known eastern Atlantic species of both the genus and family and has morphology that is, for the most part, typical of the genus. Other species currently assigned to Mithrax but not included in our molecular or morphological analyses also remain treated provisionally within Mithrax (see Systematic Account below). While Mithrax is topologically positioned nearest Teleophrys Stimpson, 1860 in our tree, support is lacking (<50/57). The two species of Teleophrys are, however, closely related to Mithraculus ruber Stimpson, 1871 with strong support (95/100), which underpins the reassignment of M. ruber to Teleophrys (Fig. ...
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... analysis of molecular phylogenetic relationships within Mithracidae s.s., 72 ingroup taxa representing 9 genera and 36 species were included, along with two species of the genus Libinia Leach, 1815 as outgroup taxa (Fig. 2). Our final alignment for this analysis of mitochondrial genes included a total of 1460 bp, these being the sum of 422 bp for 16S, 378 bp for 12S, and 658 bp for COI sequence data (excluding primer regions). In general, all analyses produced strong support (>75% bootstrap/posterior probability) for clades that we define as genera but low support (<50% bootstrap/posterior probability) for the topological relationships among these genera (Figs 1, 2). It was also obvious that many species did not group with congeners assigned traditionally on the basis of ...
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... hispidus, M. pleuracanthus, M. spinosissimus, and M. tortugae group together (65/69) under the herein proposed Damithrax, gen. nov. (Fig. 2). Nemausa acuticornis, N. cornuta, N. spinipes, and Mithrax sinensis group together in a strongly supported clade (96/100). This separates Nemausa as a distinct genus and also provides evidence that M. sinensis belongs to this group, as evident in its treatment under Nemausa sinensis, in our tree. Strong evidence is provided for continued recognition of N. acuticornis as a distinct species, given its highly supported grouping with N. sinensis and N. spinipes (98/100), independent of N. ...
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... is represented in our analyses by three species that grouped into a well supported clade (Fig. 2). The two Atlantic species (T. aspera Rathbun, 1901 andT. puella Stimpson, 1860) show a close genetic relationship while the Pacific species (T. erosa Bell, 1836) is more distantly related. Thoe is a sister genus to the newly established genus Petramithrax in our phylogenetic tree topology, but its genetic relationship to Petramithrax, while supported (67/94), is not obvious in morphology (Fig. 3L, O). These sister genera form a clade together with Hemus, but not one that conforms to morphological characters proposed in diagnosis of the tribe Thoini Štev ci c, 1994 (see Štev ci c 2005). A relationship between Thoe and Hemus is somewhat suggested by morphological (Fig. 3C, O) Fig. 2) it is allied with Mithrax, albeit with low support (<50/57). The varied development of cristae or lamelliform lobes on pereopods in some species of Teleophrys likely represents convergence with character states in Hemus and Thoe, as all three tend to occupy similar niches in tropical epifaunal fouling assemblages. The carpal segments of the pereopods in Teleophrys are, however, more rectangular and less dilated than in either Hemus or Thoe (Fig. 3C, N, O). The close genetic relationship between Teleophrys and Mithraculus ruber leads us to transfer M. ruber to Teleophrys. Thus, we cannot agree with the positioning of this species as inferred in the single gene analysis of Baeza et al. (2010: under misspelling 'Mithraculus rubber'). Species of Teleophrys have the fused basal antennal segment somewhat narrowed anteriorly, its margin never armed with more than 1 or 2 blunt teeth or tubercles, and the orbital margin overall very weakly armed; this combination of characters sets the genus apart from almost all species of other mithracid genera (see Teleophrys, Systematic ...
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... is represented in our analyses by three species that grouped into a well supported clade (Fig. 2). The two Atlantic species (T. aspera Rathbun, 1901 andT. puella Stimpson, 1860) show a close genetic relationship while the Pacific species (T. erosa Bell, 1836) is more distantly related. Thoe is a sister genus to the newly established genus Petramithrax in our phylogenetic tree topology, but its genetic relationship to Petramithrax, while supported (67/94), is not obvious in morphology (Fig. 3L, O). These sister genera form a clade together with Hemus, but not one that conforms to morphological characters proposed in diagnosis of the tribe Thoini Štev ci c, 1994 (see Štev ci c 2005). A relationship between Thoe and Hemus is somewhat suggested by morphological (Fig. 3C, O) Fig. 2) it is allied with Mithrax, albeit with low support (<50/57). The varied development of cristae or lamelliform lobes on pereopods in some species of Teleophrys likely represents convergence with character states in Hemus and Thoe, as all three tend to occupy similar niches in tropical epifaunal fouling assemblages. The carpal segments of the pereopods in Teleophrys are, however, more rectangular and less dilated than in either Hemus or Thoe (Fig. 3C, N, O). The close genetic relationship between Teleophrys and Mithraculus ruber leads us to transfer M. ruber to Teleophrys. Thus, we cannot agree with the positioning of this species as inferred in the single gene analysis of Baeza et al. (2010: under misspelling 'Mithraculus rubber'). Species of Teleophrys have the fused basal antennal segment somewhat narrowed anteriorly, its margin never armed with more than 1 or 2 blunt teeth or tubercles, and the orbital margin overall very weakly armed; this combination of characters sets the genus apart from almost all species of other mithracid genera (see Teleophrys, Systematic ...
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... analysis groups most recently assigned members of the much-debated Mithraculus into a well supported clade (100/100) ( Fig. 2), notably excluding its previous member, Teleophrys ruber (Stimpson, 1871). In all iterations of the analysis, this clade appeared to be distinct and was never recovered as sister to either Mithrax s.s. or Mithrax ...
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... (sensu Ng et al. 2008) is represented in our molecular analyses by three species that do not form a monophyletic group (Fig. 2). Most notably, the Pacific species Microphrys branchialis Rathbun, 1898 is positioned far from the Atlantic taxa as a sister to the clade encompassing Damithrax, Nemausa, Ala, and Nonala, albeit with only moderate support (72/94). Herein removed from Microphrys s.s., the Atlantic species that we now treat as Omalacantha antillensis (Rathbun, 1901) and O. bicornuta (Latreille, 1825) are grouped together with strong support (99/100), but support is lacking for their position as a sister to Teleophrys + Mithrax (<50/57). While the type species of Microphrys, M. weddelli H. Milne Edwards, 1851, was not included in the molecular analyses, morphological examinations of O. antillensis, O. bicornuta, O. interrupta (Rathbun, 1920), M. platysoma (Stimpson, 1860), M. triangulatus (Lockington, 1877), and M. weddelli reveal minor but consistent differences between the Atlantic clade (comprising O. antillensis and O. bicornuta) and putative congeners with an eastern Pacific distribution. This is, for example, true in the shape of the third maxilliped merus at the insertion of the palp, which is consistently of one form for Pacific species and another for Atlantic (see Microphrys, Systematic Account). Morphology thus underpins our provisional restriction of Pacific species to the genus Microphrys s.s. and removal of Atlantic species to the resurrected genus Omalacantha Streets, 1871 (see Systematic ...

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... Sequences obtained by this study for A. aculeatus and A. verrucosus were combined with those from Baeza et al. (2010), Windsor and Felder (2014) and Assugeni et al. (2016) which are available on GenBank. Locality information and GenBank accession numbers for all taxa included in the molecular analysis are provided in Table 1. ...
... Amphithrax verrucosus is well supported (ML = 91%, Pp = 0.989) as a separate species and sister taxa to A. aculeatus. All specimens previously identified by Windsor and Felder (2014) as A. aculeatus (ULLZ 9148 and 4534) and an additional specimen from the ULLZ collection (ULLZ 13596) were reidentified as A. verrucosus based on the morphological examination and the sequences from Barbados (see supplementary material). The phylogenetic tree shows divergence of sequences from ULLZ (4534, 9148, 13596) and MO-BR-C-1529. ...
... In the same work, Herbst (1790) described Cancer spinipes, and both its description and figure resemble A. verrucosus. However, Windsor and Felder (2014) selected the lectotype of Cancer aculeatus Herbst, 1790 as the neotype for Cancer spinipes Herbst, 1790, establishing these names as objective synonyms. Due to lack of material for Cancer spinipes Herbst, 1790, we have opted to maintain its status as a synonym of A. aculeatus. ...
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... Few published studies include DNA sequences of Pisa species. Windsor and Felder (2014) provided sequences for the 12S, 16S, 18S and COI genes for a single specimen of P. tetraodon. Raupach et al. (2015) gave two different COI sequences for P. armata, and Mugnai et al. (2023) provided two identical COI sequences for P. nodipes, although one is submitted under the name of P. armata. ...
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... However, most of the deep relationships that we do observe, even with poor nodal support from ML, are congruent with the broad results retrieved from target capture of over 400 loci (Wolfe et al. 2019), except for the position of Menippidae (stone crabs). Our topologies for the relationships within superfamilies are largely congruent with previous Sanger data , with some families exchanging places (e.g., Evans 2018; Mendoza et al. 2022) and some other groups with previous conflict still unresolved (Hultgren and Stachowicz 2008;Lai et al. 2014;Windsor and Felder 2014). The position of Dorippoidea nested well within heterotremes is consistent with previous molecular analysis ), but may confound hypotheses of early fossils that assume dorippoids are the earliest eubrachyuran branch (e.g., Guinot et al. 2019). ...
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... In the synonymy list, only the most important taxonomic accounts for each species are included (i.e., not an exhaustive list for each species). The opinions of Guinot (1967), Guinot & Tavares (2003), Manning & Chace (1990) and Windsor & Felder (2014, 2017 are mostly followed. ...
... Rathbun (1921: 83) reported one male specimen of Mithrax pilosus that was collected by the Barbados-Antigua Expedition in 1918 as a first record for Barbados (USNM 1519303); however, this species was previously reported by Schomburgk (1848: 65). Jones (1969: 380) Windsor & Felder (2017) corrected some points presented in Windsor & Felder (2014) regarding the taxonomy of Mithrax Latreille, 1816 and established the genus Amphithrax Windsor & Felder, 2017 to accommodate several species previously included in Mithrax. We do not follow the synonym between Amphithrax aculeatus and Amphithrax verrucosus; instead, we follow the opinions of Ng et al. (2008: 123), Carmona-Suárez & Poupin (2016: 372, figs. ...
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... The new genus possesses all these characteristics and is therefore referred to this subfamily. The new genus Eoparanaxia has similarities with Paranaxia, in the general outline of the carapace, showing a very elongated pseudorostral spines, parallel and distally bifid, slightly differentiated dorsal regions, with aligned spines in the dorsal part of the mesobranchial region, and well developed mesobranchial and intestinal spines (i.e., Windsor and Felder, 2014), sometimes beyond posterior carapace margin (see Rathbun, 1924, andHara, 2016, p. 128, figure 2; p. 129, figure 3). ...
... Recientemente, la familia Mithracidae ha sido ampliamente analizada a partir de caracteres morfológicos e identificación molecular, con el propósito de la reubicación de géneros y establecimiento de nuevas relaciones filogenéticas (Windsor y Felder, 2009, 2014, 2017 (Windsor & Felder, 2014); Omalacantha (Streets, 1871) y Petramithrax (Windsor & Felder, 2014). Posteriormente, Klompmaker et al. (2015) añadieron el género Maguimithrax, anteriormente renombrado por Windsor y Felder (2014) como Damithrax. ...
... Recientemente, la familia Mithracidae ha sido ampliamente analizada a partir de caracteres morfológicos e identificación molecular, con el propósito de la reubicación de géneros y establecimiento de nuevas relaciones filogenéticas (Windsor y Felder, 2009, 2014, 2017 (Windsor & Felder, 2014); Omalacantha (Streets, 1871) y Petramithrax (Windsor & Felder, 2014). Posteriormente, Klompmaker et al. (2015) añadieron el género Maguimithrax, anteriormente renombrado por Windsor y Felder (2014) como Damithrax. ...
... Recientemente, la familia Mithracidae ha sido ampliamente analizada a partir de caracteres morfológicos e identificación molecular, con el propósito de la reubicación de géneros y establecimiento de nuevas relaciones filogenéticas (Windsor y Felder, 2009, 2014, 2017 (Windsor & Felder, 2014); Omalacantha (Streets, 1871) y Petramithrax (Windsor & Felder, 2014). Posteriormente, Klompmaker et al. (2015) añadieron el género Maguimithrax, anteriormente renombrado por Windsor y Felder (2014) como Damithrax. ...
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... 1). Molecular data analysis Sequences generated for this study were combined with those from Windsor and Felder (2014;2017) and other sequences available from GenBank. Locality information and GenBank accession numbers for taxa included in the molecular analyses are provided in Tab. 1. ...
... The clade composed of Petramithrax pygmaeus (Bell, 1836a) and Hemus has low support (40 %) as a sister group to Thoe (Fig. 1). This could be a result of the different sequences used by Windsor and Felder (2014) and the present study. All three genera include very small crabs that inhabit subtidal reefs and are commonly associated with Pocillopora Lamarck, 1816 and other corals, sponges, shell hash, and reef rubble (Rathbun, 1925;Garth, 1958). ...
... Streets and Kingsley (1877: 104) examined the specimens and attested them as "plainly" T. sulcata. While reviewing this species, Windsor and Felder (2014) observed morphological variability among specimens forming a morphological continuum between species and subspecies. The review and the comparison of the distinguishing morphological characters of T. erosa, T. panamensis, and T. sulcata proposed by Garth (1958: 431-432), along with his notes, led Windsor and Felder (2014) to synonymize all Pacific species. ...
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Thoe Bell, 1836 is amphi-American in distribution and includes small spider crabs with cryptic habits and rare records in the literature and carcinological collections. The taxonomy of the three recent and two fossil species currently known in the genus has been revised. A phylogenetic reconstruction conducted here based on the mitochondrial markers COI and 16S rRNA supports the monophyletic status of Thoe. The resulting phylogeny suggests a western Atlantic to eastern Pacific radiation of the genus.
... The brachyuran family Mithracidae MacLeay, 1838 belongs to the superfamily Majoidea Samouelle, 1819, which includes species usually referred to as spider crabs. This family encompasses 14 genera and 60 species (Windsor & Felder 2014WoRMS 2021), which exhibit remarkable disparity in terms of body size, coloration, morphology, and behavior (Rathbun 1925;Baeza et al. 2010). However, the taxonomy accepted for members of the family Mithracidae has changed several times in recent decades and remains unstable at the level of genera (Windsor & Felder 2014. ...
... This family encompasses 14 genera and 60 species (Windsor & Felder 2014WoRMS 2021), which exhibit remarkable disparity in terms of body size, coloration, morphology, and behavior (Rathbun 1925;Baeza et al. 2010). However, the taxonomy accepted for members of the family Mithracidae has changed several times in recent decades and remains unstable at the level of genera (Windsor & Felder 2014. Therefore, several studies have been carried out with substantial proposals to the taxonomy of this family, based on morphological adult and larval characteristic studies, molecular phylogenetic evidence, in addition to spermatozoa and spermatophore ultrastructure data (Rhyne et al. 2006;Windsor & Felder 2009;Baeza et al. 2010;Klompmaker et al. 2015;Windsor & Felder 2014Assugeni et al. 2017). ...
... However, the taxonomy accepted for members of the family Mithracidae has changed several times in recent decades and remains unstable at the level of genera (Windsor & Felder 2014. Therefore, several studies have been carried out with substantial proposals to the taxonomy of this family, based on morphological adult and larval characteristic studies, molecular phylogenetic evidence, in addition to spermatozoa and spermatophore ultrastructure data (Rhyne et al. 2006;Windsor & Felder 2009;Baeza et al. 2010;Klompmaker et al. 2015;Windsor & Felder 2014Assugeni et al. 2017). ...
Article
The complete larval development of Amphithrax hemphilli was described, illustrated, and compared with that of the previously described larvae of the genus Mithrax sensu lato. Specimens of A. hemphilli were sampled from the northeastern Brazilian coast. The larval development of A. hemphilli consisted of two zoeal stages and one megalopa. Amphithrax hemphilli shows morphological features in all stages of larval development that differ from those observed in other species of Amphithrax, Mithrax, and Maguimithrax. In the first larval stage, A. hemphilli was the only species with two aesthetascs on the antennule and the coxal endite of the maxillule with five setae. In the second larval stage, A. hemphilli was distinguished from the other species of the genus Mithrax sensu lato by the presence of five setae on the basial endite of the maxilla and 26 setae in the scaphognathite of the maxilla. Finally, the megalopa stage of A. hemphilli is characterized by the number of aesthetascs on the three-segmented exopod of the antennule, the setation of both the endopod of the third maxilliped, and pleonites 1–6. Based on the results obtained in the present study, we verified that the more advanced the larval stage, the more distinct characteristics are observed among species of the genus Mithrax sensu lato, facilitating the identification of species through later larval stages.
... Mithraculus is amphi-American in distribution and is currently formed by nine species: Mithraculus cancasense Türkay, 1967) (Herbst, 1801) [Cancer] (WA); M. denticulatus (Bell, 1835) [Mithrax] (EP); M. forceps A. Milne-Edwards, 1875 (WA); M. nodosus (Bell, 1835) [Mithrax] (EP); M. rostratus (Bell, 1835) [Mithrax] (EP); and M. sculptus (Lamarck, 1818 [Maia] (WA). Mithraculus ruber Stimpson, 1871a (WA), has been transferred to Teleophrys (see below) by Windsor & Felder (2014). (Melo 1996;Serejo et al. 2006;Bouzon & Freire 2007;Felder et al. 2009). ...
... Remarks. The resurrection of the genus Omalacantha from the synonymy of Microphrys was underpinned by molecular data combined with less obvious morphological support (Windsor & Felder 2014). Clearly, the morphological differences between the two genera need to be more intensively investigated. ...
... Omalacantha as understood by Windsor & Felder (2014) consists of four species, namely: O. antillensis (Rathbun, 1920) [Microphrys], type locality: off Montego Bay Point, Jamaica; O. bicornuta (Latreille, 1825) [Pisa], type locality: "Nouvelle Hollande"-could be either Dutch Guiana, Saba, Saint Eustace, St. Martin's or Tobago, at that time under Dutch control (Boone 1930a: 111); O. garthi (Lemos de Castro, 1953) [Eucinetops], type locality: Barra de Guaratiba, state of Rio de Janeiro; and O. interrupta (Rathbum, 1920) [Microphrys]. ...
Article
Forty species of shore and shallow water brachyuran crabs are reported from the remote oceanic Archipelago Trindade and Martin Vaz (TMV), 11 of which were previously known from Trindade Island, 28 are recorded from the Archipelago for the first time, and one is a new species, Epialtus parvulus sp. nov. This brings the total known shore and shallow water decapod fauna of TMV to 99 species. The opportunity is taken here to elaborate on the taxonomy of the species reported from TMV. One new genus, Mecataleptodius, is recognized for two species, Cancer parvulus Fabricius, 1793, its type species, and Cataleptodius olsoni Manning & Chace, 1990, endemic to Ascension Island. The following seven nominal species are synonymized: Acanthonyx dissimulatus Coelho in Coelho & Torres, 1993; A. scutiformis (Dana, 1851a); Dromia gouveai Melo & Campos Junior, 1999; Epialtus portoricensis Rathbun, 1923; Ranilia guinotae Melo & Campos Junior, 1994; R. saldanhai H. Rodrigues da Costa, 1970; and Omalacantha garthi (Lemos de Castro, 1953). TMV is a subset of the western Atlantic crab fauna (23 species or 57.5%), also including 8 (20%) amphi-Atlantic species, 3 (7.5%) amphi-American, 3 (7.5%) endemic, 2 (5%) cosmopolitan, 1 (2.5%) known from both sides of the Atlantic and the Indo-West Pacific. The crab fauna of TMV, Ascension (AS) and Saint Helena (SH) are compared with one another as well as with that of the oceanic islands Fernando de Noronha (FN) and the Rocas Atoll (RA) by means of nonmetric multidimensional scaling (NMDS) applied to a dissimilarity matrix generated by Jaccard’s coefficient. TMV-FN-RA and AS-SH clustered into two distinct groups according to the composition of their crab fauna. The NMDS analysis ranked the species that cause segregation of the crab faunal assemblages among TMV, AS, and SH, while revealing a gradient in species composition between the two groups of islands (TMV-FN-RA and AS-SH) formed by the amphi-Atlantic species. Island isolation, age and size alone do not explain the existing differences in species richness among TMV, AS and SH. The taxonomic composition of the shallow water fauna in these islands is likely to have been determined by several variables that were interwoven differently along their evolutionary history.