FIG 7 - uploaded by Rodrigo Monjaraz-Ruedas
Content may be subject to copyright.
Short-tailed whipscorpions (Schizomida: Hubbardiidae Cook, 1899), pedipalps, retrolateral view. A. Ambulantactus aquismon, sp. nov., ♂ (CNAN T1294). B. Ambulantactus montielae, sp. nov., ♂ (CNAN T1291). C. Baalrog magico (Monjaraz-Ruedas and Francke, 2018), comb. nov., ♂ (CNAN T1163). D. Harveyus contrerasi, sp. nov., ♂ (CNAN T1276). E. Heteroschizomus kekchi, sp. nov., ♂ (CNAN T1280). F. Nahual bokmai, sp. nov., ♂ (CNAN T1282). G. Schizophyxia lukensi (Rowland, 1973), comb. nov., ♂ (CNAN Sz25). H. Stenochrus gruta Monjaraz-Ruedas and Francke, 2018, ♂ (CNAN T1155). I. Troglostenochrus valdezi (Monjaraz-Ruedas, 2012), comb. nov., ♂ (CNAN T698). Scale bars = 0.2 mm.
Contexts in source publication
Context 1
... the apical process may also be interpreted as a rounded margin, i.e., not projected. The terms applied to the states of this character follow botanical terminology for leaf tips: "acute" is the most common shape observed in North American schizomids (fig. 7E, C); "acuminate" is the alternate form of the state with the margin extremely projected ( fig. 7F, I); "obtuse triangle" has the margin not projected with a wide internal angle (fig. 7D); "bump" or "tubercle" is observed mainly among heteromorphic males (fig. 7H); digitiform is common in Surazomus and some species of ...
Context 2
... Fe 5 , Fv 1 , Fv 2 , Ter 7 and Tir 5 acuminate, setiform; pygidial flagellum (♂) ovate, subrhomboidal, or bulbous (figs. 16G-I, 23D-F), with pair of dorsosubmedian pits or depressions not fused posteriorly; spermathecae (♀), lateral lobes slender, filiform (figs. 8B, D)........................12 12. Pedipalp trochanter, apical process with bump ( fig. 7C, H); pygidial flagellum (♂) subrhomboidal, and acuminate or lanceolate posteriorly ( fig. 19A-C), with pair of shallow dorsosubmedian pits; spermathecae (♀), lateral lobes at least 2/3 the length of, or longer than, median lobes, and with apical bulbs ( fig. 8C, ...
Context 3
... gen. nov. -Pedipalp trochanter, apical process acute or absent (fig. 7B); pygidial flagellum (♂) elliptical, cordate or bulbose, and not acuminate or lanceolate posteriorly ( fig. 23D-F), with pair of deep dorsosubmedian pits or without dorsal depressions; spermathecae (♀), lateral lobes at most half the length of median lobes, and without apical bulbs (figs. 8B, 9E).....................................13 ...
Context 4
... lateral lobes of the female spermathecae, mistakenly reported as absent by Rowland and Reddell (1980), are extremely reduced (fig. 8B). The discovery of S. magico revealed the unique bulbous pygidial flagellum of the male. The lateral lobes of the spermathecae are unusually variable in size in this species (Monjaraz-Ruedas and Francke, 2018: 210, fig. 70). These characters are diagnostic for Baalrog, gen. ...
Context 5
... Pedipalps homeomorphic (fig. 7D); 1.73× (♂) or 1.59× (♀) longer than propeltidium. Trochanter with small rounded apical process; prolateral surface with small apical spur. Femur 1.94× longer than high; retroventral margin with setae Fe 1 , Fe 5 , Fev 1 , and Fev 2 acuminate; prolateral surface with row of three ventral acuminate setae (Fmv [1][2][3] ...
Context 6
... Pedipalps homeomorphic (fig. 7E); 1.66× (♂) or 1.69× (♀) longer than propeltid-ium. Trochanter with small acute apical process; prolateral surface with small medial spur. Femur 1.8× longer than high; retroventral margin with setae Fe 1 , Fe 5 , Fev 1 , and Fev 2 acuminate; prolateral surface with row of three ventral setae (Fmv [1][2][3] ) and two dorsal setae (Fmd 2 ...
Context 7
... (except in N. bokmai, sp. nov.) ( fig. 13F). Propeltidium anterior process with two anterior setae (one posterior to the other) and three pairs of dorsosubmedian setae (fig. 11A); corneate eyes absent. Metapeltidium entire. Tegument without clavate setae. Pedipalps homeomorphic; trochanter with mesal spur and projected, fan-shaped apical process (fig. 7F); femur Fv 1 and Fv 2 setae spiniform, Fvr 1-3 setae present (seta Fvr 4 present in N. lanceolatus, comb. nov., polymorphic in N. bokmai and N. pallidus, comb. nov.); patella with four or five acuminate Pe setae and five feathered Pm setae; tibial setal formula 5(6)-5-6 (Ter-Tmr-Tir) ( fig. 14C). Leg IV femur anterodorsal margin ...
Context 8
... and differs from the enlarged flagella observed in species of Heteroschizomus, stat. rev., by the absence of a posterior constriction. Diagnosis: Nahual bokmai, sp. nov., may be separated from other species of Nahual, gen. nov., by the following characters: the pedipalp trochanter of the male bears a long, broad, and triangular apical process (fig. 7F); the male pygidial flagellum is elliptical with the posterior half acuminate ( fig. 19G-I); the lateral tips of the chitinized arch of the female spermathecae ( fig. 9A) are extremely widened; and the median and lateral lobes of the spermathecae are linear, with the lateral lobes slightly shorter than the median ...
Context 9
... Pedipalps homeomorphic (fig. 7F); 2.16× (♂) or 1.7× (♀) longer than propeltidium. Trochanter with broad triangular apical process; prolateral surface with long medial spur. Femur 2.1× longer than high; retroventral margin with setae Fe 1 and Fe 5 acuminate, Fev 1 and Fev 2 spiniform; prolateral surface with row of three ventral spiniform setae (Fmv 1-3 ) and two ...
Context 10
... with G3-3 setae situated anteriorly ( fig. 13G). Propeltidium anterior process with two anterior setae (one posterior to the other) and three pairs of dorsosubmedian setae (fig. 11A); corneate eyes absent. Metapeltidium entire. Tegument without clavate setae. Pedipalps homeomorphic; trochanter with mesal spur, apical process acute, not projected (fig. 7G); femur Fv 1 and Fv 2 setae acuminate, Fvr 1-3 setae present; patella with four acuminate Pe setae and four feathered Pm setae; tibial setal formula 3-3-4 (TerTmr-Tir) ( fig. 14D). Leg IV femur anterodorsal margin produced at ca. 90° angle. Opisthosomal tergite II with one pair of setae (Dm). Opisthosomal segments IX-XII not elongated; ...
Context 11
... absent. Metapeltidium entire. Tegument without clavate setae. Pedipalps heteromorphic (as in Stenochrus chimalapas Stenochrus gruta Monjaraz-Ruedas and, with femur, patella and tibia elongated; trochanter with small mesal spur and without apical process (except in S. alcalai and S. gruta) with an acute apical process and a bump, respectively) (fig. 7H); femur Fv1 and Fv 2 setae acuminate (except in S. pecki, with spiniform setae), Fvr 1-3 setae present; patella with three acuminate Pe setae and three or four feathered Pm setae; tibial setal formula 3-3-4 (Ter-Tmr-Tir) (except S. alcalai with 3-3-5) ( fig. 14D). Leg IV femur anterodorsal margin produced at ca. 90° angle. Opisthosomal ...
Similar publications
The genus Caerulea FORSTER, 1938 is reviewed, with C. guizhouensis spec. nov. described from W. Guizhou.
Citations
... The Order Schizomida (Arachnida: Tetrapulmonata) -the shorttailed whipscorpions -currently includes about 345 extant plus 16 fossil species (Monjaraz-Ruedas et al., 2019Giribet & Moreno-González, 2021). They have a worldwide distribution mostly in tropical and subtropical areas (Reddell & Cokendolpher, 1995). ...
Herein we report for the first time a schizomid for the Southern region of Brazil, Stenochrus portoricensis Chamberlin, 1922 (Schizomida: Hubbardiidae), found in association with termite nests. This is the southernmost record of any schizomid for the Neotropical region. We hypothesize that the species was recently introduced by the sudden population growth of Florianópolis – along with the intense touristic activity – which might have contributed to the inadvertent transportation of this species.
... After that, however, most studies continued with the flagellum-spermathecae-only approach, but including newer technologies, such as digital photographs. SEM images have been used since only in a series of studies (chelicera: [22][23][24][25][26]; chelicera and male flagellum: [27,28]; female flagellum: [24]). Also, Giupponi et al. [29] and Villarreal et al. [30] published SEM illustrations of the chelicera, pedipalp, male and female flagella and some leg details. ...
... sexual dimorphism in the chelicera of S. algodoal. The few G5B setae observed on the female chelicera (three setae in the female; eight in the male; Table 2) may be a variation, since females of other species, such as Surazomus cuenca (Rowland & Reddell, 1979) and Surazomus chiapasensis seem to have about eight G5B setae [28,30], as males. That leads us to conclude that the sexual selective pressure may be affecting the counterpart-the male pygidial flagellum. ...
... Despite the diversity of setae and other structures in the chelicera of a single species, their contribution to phylogenetic reconstruction within Hubbardiines seems of limited use, since all the seta types seem to be present and similar in number and form throughout the subfamily (e.g. [28]). However, these structures may serve as a morphological marker for the group. ...
The male genitalia of Surazomus algodoal Ruiz & Valente, 2017 are described. Since this is the first attempt to describe male genitalia in the schizomids, we propose new terminology to describe the sclerites of the male genital chamber in the order. We believe that the male genitalia may provide a large set of characters for taxonomic research in schizomids, as the female genitalia have proven to do. The setae and other cuticular structures were investigated using light and scanning electron microscopy (SEM). The female of S. algodoal is described for the first time, including the genitalia, and we highlight the sexual dimorphism present in the species. We also present descriptions of: 1) microtrichia, with respect to their presence/length/distribution, 2) cuticular sensilla, glandular openings and other cuticular structures, proposing new terminology, 3) setae of the pedipalpal trochanter and tarsus. Some changes on the terminology of setae of pedipalpal tibia are also given. These advances may be useful for description in other schizomids. The pygidial glands, mostly ignored in modern papers, were assessed and commented upon.
... oculiferans) show heteromorphic pedipalps, extending the evolutionary age of this character state back to the mid-Cretaceous. Heteromorphism in male pedipalps of Recent schizomids has been noted by various authors (Reddell & Cokendolpher, 1995;Harvey, 2001a;Armas & Teruel, 2002;Monjaraz-Ruedas et al., 2019). The reason for this is still unknown, but it could be driven by sexual selection. ...
Short-tailed whipscorpions are a small arachnid order with 379 described species from (sub)tropical regions of the world. The order has a fragmentary fossil record but has left a rich, albeit hitherto undocumented, fossil legacy in mid-Cretaceous amber from Myanmar, also called Burmese amber or Burmite. Here, we review a large series of schizomids from this type of amber and propose four new genera for seven new species. In addition, we include a short revision of all schizomid genera based on 14 morphological characters. Although none of the fossils can be assigned to Recent genera, they resemble the modern fauna in general morphology and highlight relative morphological stasis in this arachnid order. The fossils show a general trend towards eye reduction, but some of the specimens retain corneate eyes. Comparative data suggest affinities of the fossils with the Recent Old World fauna rather than American relatives. Furthermore, the fossils provide evidence for a Gondwanan origin of the Burma Terrane. Overall, the data suggest a high diversity at both species and genus levels for Burmese amber schizomids. We discuss potential reasons for this richness that will probably necessitate the description of additional genera and species in future studies.
... The male flagellar shape is phenotypically diverse and characterizing them for quantitative analysis has proven difficult. For example, linear morphometric ratios 20 and discrete states 21 have been used to code them for cladistic analyses, although, doing so reduces the available shape diversity. The male flagellum is critical for classification and species identification, and is likely important for sexual selection and reproduction, yet, there has been little research done that examines the shape diversity across schizomids 4,18 . ...
The arachnid order Schizomida is a relatively understudied group of soil-dwelling predators found on all continents except Antarctica. While efforts to understand their biology are growing, there is still much to know about them. A curious aspect of their morphology is the male flagellum, a sexually dimorphic, tail-like structure which differs in shape across the order and functions in their courtship rituals. The flagellar shape is important for taxonomic classification, yet few efforts have been made to examine shape diversity across the group. Using elliptical Fourier analysis, a type of geometric morphometrics based on shape outline, we quantified shape differences across a combined nearly 550 outlines in the dorsal and lateral views, categorizing them based on genus, family, biogeographic realm, and habitat, with special emphasis on Caribbean and Cuban fauna. We tested for allometric relationships, differences in disparity based on locations and sizes in morphospace among these categories, and for clusters of shapes in morphospace. We found multiple differences in all categories despite apparent overlaps in morphospace, evolutionary allometry, and evidence for discrete clusters in some flagellum shapes. This study can serve as a foundation for further study on the evolution, diversification, and taxonomic utility of the male flagellum.
... The present study evaluates the phylogeography of S. portoricensis and its close relatives (Monjaraz-Ruedas et al. 2019 using multilocus data from two nuclear and two mitochondrial markers for the broadest geographical sample currently available. Genetic clustering and tree-based approaches are used to evaluate genetic structure and haplotype diversity among populations and to investigate species limits. ...
Whereas morphology remains a powerful tool for the diagnosis and description of short-tailed whip scorpions, or schizomids (Order Schizomida Petrunkevitch, 1945), especially when adults of both sexes are available, the systematics of some schizomid taxa is difficult to resolve due to a lack of characters in these morphologically conserved arachnids. Stenochrus portoricensis Chamberlin, 1922, defined on a single character of the female spermathecae, is the most widespread schizomid in the New World. Numerous records in the Neotropics, from the southern United States to Brazil, throughout the Caribbean, and further afield, including the Galapagos Islands and Europe, raise the question as to whether S. portoricensis is indeed a single widespread species or a complex of multiple species with conserved morphology? The present study uses a multilocus dataset and the broadest geographical sample currently available to address the phylogeography of S. portoricensis with molecular divergence dating and ancestral area reconstruction of all currently known species of Stenochrus Chamberlin, 1922. Analyses recovered S. portoricensis as paraphyletic. Two species previously synonymized are revalidated and transferred to Stenochrus. Population structure analyses recovered the remaining samples of S. portoricensis as a single monophyletic species with low genetic divergence and comprising two subclades. Ancestral area reconstruction suggests a Mesoamerican origin for Stenochrus, which contains a widespread species, recently introduced to multiple localities. Introductions to Europe and the Caribbean occurred from a single clade in the Yucatán Peninsula, Mexico, within which genetic divergence is minimal, confirming the hypothesis of multiple independent introductions with successful colonization facilitated by parthenogenetic reproduction.
... Short-tailed whip scorpions are represented in the cave system by a single species Baalrog magico (Monjaraz-Ruedas and Francke, 2018), which was originally described under the genus Stenochrus Chamberlin, 1922 and then transferred to the newly described genus Baalrog Monjaraz-Ruedas, Prendini and Francke, 2019, which harbors species restricted to the cave environment. Other members of genus Baalrog are distributed in cave systems in Valle Nacional, Oaxaca, and Atoyac, Veracruz [30]. Although there are other undescribed species of schizomids in the Sierra Mazateca outside the caves, those belong to different genera and are apparently not related to the species distributed inside Sistema Huautla [31]. ...
Sistema Huautla is the deepest cave system in the Americas at 1560 m and the fifth longest in Mexico at 89,000 m, and it is a mostly vertical network of interconnected passages. The surface landscape is rugged, ranging from 3500 to 2500 masl, intersected by streams and deep gorges. There are numerous dolinas, from hundreds to tens of meters in width and depth. The weather is basically temperate subhumid with summer rains. The average yearly rainfall is approximately 2500 mm, with a monthly average of 35 mm for the driest times of the year and up to 500 mm for the wettest month. All these conditions play an important role for achieving the highest terrestrial troglobite diversity in Mexico, containing a total of 35 species, of which 27 are possible troglobites (16 described), including numerous arachnids, millipedes, springtails, silverfish, and a single described species of beetles. With those numbers, Sistema Huautla is one of the richest cave systems in the world.
... The order Schizomida Petrunkevitch, 1945, commonly known as the short-tailed whipscorpions, schizomids or tartarids, is among the less diverse, or minor, arachnid orders (Harvey, 2003). It currently includes two families: the Protoschizomidae Rowland, 1975, endemic to North America, comprising two extant and one fossil genera with 16 species, and the Hubbardiidae Cook, 1899, with 65 extant genera and 345 species worldwide (Harvey, 2003(Harvey, , 2013Monjaraz-Ruedas et al., 2019). There are 33 genera of Hubbardiidae (Reddell & Cokendolpher, 1995;Harvey, 2013;Monjaraz-Ruedas et al., 2019) in the New World. ...
... It currently includes two families: the Protoschizomidae Rowland, 1975, endemic to North America, comprising two extant and one fossil genera with 16 species, and the Hubbardiidae Cook, 1899, with 65 extant genera and 345 species worldwide (Harvey, 2003(Harvey, , 2013Monjaraz-Ruedas et al., 2019). There are 33 genera of Hubbardiidae (Reddell & Cokendolpher, 1995;Harvey, 2013;Monjaraz-Ruedas et al., 2019) in the New World. ...
... Given the paucity of phylogenetic analyses on schizomids, it is unsurprising that the monophyly and phylogenetic relationships of Stenochrus sensu Reddell & Cokendolpher (1991, 1995 had never been tested, beyond the unpublished analysis of Rowland (1975), and the inclusion of exemplar species in recent analyses of schizomid phylogeny based on morphology Rowland & Reddell (1979a, b, 1980, 1981. Stenochrus was also paraphyletic in the molecular analysis of Clouse et al. (2017), based on two nuclear and two mitochondrial gene markers for 240 samples, which included several individuals of S. portoricensis, one Stenochrus sbordonii (Brignoli, 1973) The present contribution provides the first phylogeny of Stenochrus and related genera, based on 61 morphological characters and 2991 aligned DNA nucleotides from two markers in the nuclear genome, -Ruedas et al., 2019, Heteroschizomus Rowland, 1973, Nahual Monjaraz-Ruedas et al., 2019, Schizophyxia Monjaraz-Ruedas et al., 2019, and Troglostenochrus Monjaraz-Ruedas et al., 2019 the internal transcribed spacer (ITS) and 28S rDNA, and two markers in the mitochondrial genome, 12S rDNA and cytochome c oxidase subunit I (COI), for a comprehensive sample of 73 taxa. Separate and simultaneous analyses of the morphological and molecular data sets were conducted with different optimality criteria and analytical parameters: Bayesian Inference, Maximum Likelihood and parsimony with equal and implied weighting. ...
Until recently, the Nearctic short-tailed whipscorpion genus, StenochrusChamberlin, 1922, included 27 species distributed primarily in Mexico, the USA and Central America. Morphological disparity among its species, associated with their adaptation to diverse habitats, raised the question as to whether Stenochrus was monophyletic. The phylogenetic relationships among short-tailed whipscorpions have only recently begun to be explored, and the monophyly of Stenochrus had never been tested. The present contribution provides the first phylogeny of Stenochrus and related genera, based on 61 morphological characters and 2991 aligned DNA nucleotides from two nuclear and two mitochondrial gene markers, for 73 terminal taxa. Separate and simultaneous analyses of the morphological and molecular data sets were conducted with Bayesian Inference, Maximum Likelihood, and parsimony with equal and implied weighting. Terminals represented only by morphological data (‘orphans’) were included in some analyses for evaluation of their phylogenetic positions. As previously defined, Stenochrus sensuReddell & Cokendolpher (1991, 1995) was consistently polyphyletic and comprised eight monophyletic clades, justifying its reclassification into eight genera including Heteroschizomus Rowland, 1973, revalidated from synonymy with Stenochrus by Monjaraz-Ruedas et al. (2019). Rowland & Reddell’s (1980)mexicanus and pecki species groups were consistently paraphyletic. Orphans grouped with the most morphologically similar taxa.
Despite a global distribution throughout the tropics and sub-tropics, the order Schizomida (Arachnida) is heavily understudied and the phylogeny of the group is poorly understood. Identification keys are only available for some regions or genera but not for the entire order. (1) comprehensively reviewed the entire schizomid fauna and established a suite of characters to define all genera known at this time. This suite of characters still depicts the foundation of modern descriptions, supplemented by recently established characters, most of them documenting setation patterns on pedipalps, flagellum and chelicerae. In this paper, we present the Schizomida Trait Data Base (STDB) containing data for 25 characters based on the entire body of schizomid literature. Characters were chosen based on their use for modern taxonomic description and availability of the data. The STDB is a powerful tool that can be used by both amateurs and experienced researchers to categorise newly found specimens, both extant and fossil, down to genus level easily. Analysis using the new database gives insight into biogeographical patterns of characters. Furthermore, we are describing a new species, †Annazomus jamesi, a fossil specimen from Burmese (Kachin) amber and investigate a small collection of extant schizomids from Ecuador, which includes the previously unrecorded female of Surazomus palenque, herein described for the first time. The taxonomic assignment of both specimens is based on the STDB, highlighting the utility of the new data base approach to schizomid systematics.
Arachnids, biogeography, Cretaceous, data base, fossil, new species, palaeontology, Schizomida, statistics, taxonomy
