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Asian forest scorpions (Scorpionidae Latreille, 1802: Heterometrinae Simon, 1879) are distributed across South and Southeast Asia. All are fossorial, constructing burrows under stones or in open ground, in habitats differing in precipitation and vegetation cover, from rainforests and tropical deciduous forests to savanna and scrubland. The systematics of these scorpions has long been confused due to bad taxonomy and the absence of a phylogenetic framework. Although the monophyly of the group was previously confirmed as part of broader phylogenetic analyses based on exemplar species, the only quantitative analysis of species‐level variation to date was based on overall similarity. This contribution presents the first species‐level phylogenetic analysis of Asian Scorpionidae, based on 186 morphological characters and 4188 aligned base‐pairs of DNA sequence data from two nuclear and three mitochondrial loci for 132 terminals including all 41 ingroup species and four outgroup species. Simultaneous analyses of the morphological and molecular datasets with parsimony, Maximum Likelihood and Bayesian Inference provided the framework for a revised classification presented elsewhere. In order to understand how adaptation following dispersal into new habitats has driven the morphological diversification of Asian forest scorpions, species were scored for 10 characters concerning morphology and burrow architecture, which contributed to an ensemble index of adaptation to habitat aridity. Species were classified into three ecomorphotypes based on the index, and ancestral state reconstruction of ecomorphotypes performed on the phylogeny. A pattern was recovered in which lineages and species occurring in different habitats on a continuum from wet (evergreen forest) to dry (savanna, scrubland) exhibited characters presumed to be adaptive and hence responsible for driving scorpion diversification. [ABSTRACT FROM AUTHOR]

The ‘Out of India’ hypothesis is often invoked to explain patterns of distribution among Southeast Asian taxa. According to this hypothesis, Southeast Asian taxa originated in Gondwana, diverged from their Gondwanan relatives when the Indian subcontinent rifted from Gondwana in the Late Jurassic, and colonized Southeast Asia when it collided with Eurasia in the early Cenozoic. A growing body of evidence suggests these events were far more complex than previously understood, however. The first quantitative reconstruction of the biogeography of Asian forest scorpions (Scorpionidae Latreille, 1802: Heterometrinae Simon, 1879) is presented here. Divergence time estimation, ancestral range estimation, and diversification analyses are used to determine the origins, dispersal and diversification patterns of these scorpions, providing a timeline for their biogeographical history that can be summarized into four major events. (1) Heterometrinae diverged from other Scorpionidae on the African continent after the Indian subcontinent became separated in the Cretaceous. (2) Environmental stresses during the Cretaceous–Tertiary (KT) mass extinction caused range contraction, restricting one clade of Heterometrinae to refugia in southern India (the Western Ghats) and Sri Lanka (the Central Highlands). (3) Heterometrinae dispersed to Southeast Asia three times during India’s collision with Eurasia, the first dispersal event occurring as the Indian subcontinent brushed up against the western side of Sumatra, and the other two events occurring as India moved closer to Eurasia. (4) Indian Heterometrinae, confined to southern India and Sri Lanka during the KT mass extinction, recolonized the Deccan Plateau and northern India, diversifying into new, more arid habitats after environmental conditions stabilized. These hypotheses, which are congruent with the geological literature and biogeographical analyses of other taxa from South and Southeast Asia, contribute to an improved understanding of the dispersal and diversification patterns of taxa in this biodiverse and geologically complex region. [ABSTRACT FROM AUTHOR]

The genera and species of the Asian forest scorpions (Scorpionidae Latreille, 1802) are revised based on a phylogenetic analysis of 186 morphological characters and 4188 base pairs of concatenated DNA sequence from three mitochondrial loci and two nuclear loci. Revision of the Asian scorpionids required a critical reappraisal of the suprageneric classification of Scorpionidae, on the basis of which the monotypic Indian scorpionoid genus, RugodentusBastawade et al., 2005, stat. rev., and its type species, Rugodentus keralaensisBastawade et al., 2005, stat. rev., are revalidated, and subfamily Rugodentinae Bastawade et al., 2005, revalidated and elevated to the rank of family, Rugodentidae Bastawade et al., 2005, stat. nov. et stat. rev.; Heterometrinae Simon, 1879, stat. nov., and Opistophthalminae Rossi, 2016, stat. nov., are elevated to the rank of subfamily; PandinopsisVachon, 1974, stat. nov., and Pandipalpus Rossi, 2015, stat. nov., are elevated to the rank of genus, resulting in two new combinations: Pandinopsis dictator (Pocock, 1888), comb. nov., and Pandipalpus viatoris (Pocock, 1890), comb. nov.; and 10 new synonyms are presented: Pandinopsini Rossi, 2016 = Pandininae Thorell, 1876, syn. nov.; Protophthalmini Rossi, 2016 = Opistophthalminae Rossi, 2016, syn. nov.; ProtophthalmusLawrence, 1969 = Opistophthalmus C.L. Koch, 1837, syn. nov.; Pandinoides (Dunlopandinoides) Rossi, 2016 = PandinoidesFet, 2000, syn. nov.; Pandinurus (Pandicaporiaccous) Rossi, 2015 = Pandiborellius Rossi, 2015, syn. nov.; Buthus defensor C.L. Koch, 1837 = Pandinurus gregoryi (Pocock, 1896), syn. nov.; Buthus heros C.L. Koch, 1837 = Pandinurus exitialis (Pocock, 1888), syn. nov.; Pandinus loweiKovařík, 2012 = Pandipalpus viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pygmaeus Rossi, 2015 = Pandipalpus viatoris (Pocock, 1890), syn. nov.; Pandinus intermedius Borelli, 1919 = Pandinurus citernii (Borelli, 1919), syn. nov. The following revisions are implemented to the classification of the Asian forest scorpions (Heterometrinae). Three former subgenera of Heterometrus Ehrenberg, 1828 are revalidated and elevated to the rank of genus: ChersonesometrusCouzijn, 1978, stat. nov. et stat. rev.; JavanimetrusCouzijn, 1981, stat. nov. et stat. rev.; and SrilankametrusCouzijn, 1981, stat. nov. et stat. rev. One subgenus is elevated to the rank of genus: GigantometrusCouzijn, 1978, stat. nov. Two new genera and eight new species are described: Deccanometrus, gen. nov.; Sahyadrimetrus, gen. nov.; Chersonesometrus bastawadei, sp. nov.; Chersonesometrus hendersoni, sp. nov.; Chersonesometrus nathanorum, sp. nov.; Chersonesometrus shivashankari, sp. nov.; Sahyadrimetrus mathewi, gen. et sp. nov.; Sahyadrimetrus tikaderi, gen. et sp. nov.; Srilankametrus couzijni, sp. nov.; Srilankametrus pococki, sp. nov. Heterometrus sensu stricto is restricted to eight species of the nominotypical subgenus, all other species, formerly placed in Heterometrus, are transferred to appropriate genera, five species are revalidated, and two subspecies elevated to the rank of species, resulting in 28 new combinations: Chersonesometrus beccaloniae (Kovařík, 2004), comb. nov.; Chersonesometrus fulvipes (C.L. Koch, 1837), comb. nov.; Chersonesometrus madraspatensis (Pocock, 1900), comb. nov.; Chersonesometrus pelekomanus (Couzijn, 1981), comb. nov. et stat. rev.; Chersonesometrus tristis (Henderson, 1919), comb. nov.; Chersonesometrus wroughtoni (Pocock, 1899), comb. nov.; Deccanometrus bengalensis (C.L. Koch, 1841), comb. nov.; Deccanometrus latimanus (Pocock, 1894), comb. nov.; Deccanometrus liurus (Pocock, 1897), comb. nov.; Deccanometrus obscurus (Couzijn, 1981), comb. et stat. nov.; Deccanometrus phipsoni (Pocock, 1893), comb. nov.; Deccanometrus ubicki (Kovařík, 2004), comb. nov.; Deccanometrus xanthopus (Pocock, 1897), comb. nov.; Gigantometrus swammerdami (Simon, 1872), comb. nov.; Gigantometrus titanicus (Couzijn, 1981), comb. nov. et stat. rev.; Heterometrus glaucus (Thorell, 1876), comb. nov. et stat. rev.; Heterometrus laevigatus (Thorell, 1876), comb. nov. et stat. rev.; Heterometrus silenus (Simon, 1884), comb. nov. et stat. rev.; Javanimetrus cyaneus (C.L. Koch, 1836), comb. nov.; Sahyadrimetrus barberi (Pocock, 1900), comb. nov.; Sahyadrimetrus kanarensis (Pocock, 1900), comb. nov.; Sahyadrimetrus rugosus (Couzijn, 1981), comb. et stat. nov.; Sahyadrimetrus scaber (Thorell, 1876), comb. nov.; Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev.; Srilankametrus gravimanus (Pocock, 1894), comb. nov.; Srilankametrus indus (DeGeer, 1778), comb. nov.; Srilankametrus serratus (Pocock, 1900), comb. nov; Srilankametrus yaleensis (Kovařík et al., 2019), comb. nov. Twenty-seven new synonyms are presented: Scorpio leiodermaDufour, 1856 = Sahyadrimetrus scaber (Thorell, 1876), syn. nov.; Palamnaeus costimanus var. β borneensis Thorell, 1876 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Palamnaeus liophysaThorell, 1888 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Palamnaeus oatesii Pocock, 1900 = Heterometrus petersii (Thorell, 1876), syn. nov.; Palamnaeus swammerdami flavimanus Pocock, 1900 = Gigantometrus swammerdami (Simon, 1872), syn. nov.; Heterometrus liophysa var. madoerensisKopstein, 1921 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus laevifronsRoewer, 1943 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Chersonesometrus) granulomanusCouzijn, 1981 = Srilankametrus caesar (C.L. Koch, 1841), syn. nov.; Heterometrus (Heterometrus) liophysa separatusCouzijn, 1981 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Heterometrus) liophysa spartanicusCouzijn, 1981 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Heterometrus) longimanus bengkalitensisCouzijn, 1981 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Heterometrus (Heterometrus) longimanus marmoratusCouzijn, 1981 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Heterometrus (Heterometrus) petersii mindanaensisCouzijn, 1981 = Heterometrus silenus (Simon, 1884), syn. nov.; Heterometrus (Heterometrus) spinifer solitariusCouzijn, 1981 = Heterometrus spinifer (Ehrenberg, 1828), syn. nov.; Heterometrus (Srilankametrus) indus laevitensusCouzijn, 1981 = Srilankametrus indus (DeGeer, 1778), syn. nov.; Heterometrus (Heterometrus) keralaensisTikader and Bastawade, 1983 = Sahyadrimetrus rugosus (Couzijn, 1981), syn. nov.; Heterometrus cimrmaniKovařík, 2004 = Heterometrus laevigatus (Thorell, 1876), syn. nov.; Heterometrus mysorensisKovařík, 2004 = Chersonesometrus tristis (Henderson, 1919), syn. nov.; Heterometrus nepalensisKovařík, 2004 = Deccanometrus bengalensis (Pocock, 1900), syn. nov.; Heterometrus rolcikiKovařík, 2004 = Sahyadrimetrus scaber (Thorell, 1876), syn. nov.; Heterometrus sejnaiKovařík, 2004 = Javanimetrus cyaneus (C.L. Koch, 1836), syn. nov.; Heterometrus tibetanusLourenço et al., 2005 = Deccanometrus bengalensis (Pocock, 1900), syn. nov.; Heterometrus liangiZhu and Yang, 2007 = Heterometrus silenus (Simon, 1884), syn. nov.; Heterometrus telanganaensis Javed et al., 2010 = Deccanometrus xanthopus (Pocock, 1897), syn. nov.; Heterometrus atrascorpiusMirza et al., 2012 = Chersonesometrus beccaloniae (Kovařík, 2004), syn. nov.; Heterometrus minotaurusPlíšková et al., 2016 = Heterometrus laevigatus (Thorell, 1876), syn. nov.; Heterometrus bastawadei Rossi, 2016 = Rugodentus keralaensisBastawade et al., 2005, syn. nov. Another 25 synonyms by previous authors are confirmed, for a total of 51 synonyms in subfamily Heterometrinae. Revised diagnoses with comparative images, and a key and distributional atlas of the genera and species are provided, along with a summary of available data for their ecology and conservation status, where applicable. [ABSTRACT FROM AUTHOR]

[Display omitted] • The first dated phylogeny of the tribe Tyrannochthoniini. • Western Ghats Tyrannochthoniini have a Gondwanan origin. • Ancient diversification patterns predating the Plio-Pleistocene climate fluctuations. • 'Museum' model of diversification supported by declining diversification rates. • Palghat Gap of the Western Ghats not an absolute barrier for dispersals. The tropical forests of the Western Ghats (WG) of India are considered 'refugia' harbouring highly diverse and endemic taxa but these refugia are under immense anthropogenic pressure. Most phylogenetic studies have explained diversity patterns across the WG using vertebrates, however, the processes impacting the highly endemic invertebrate fauna are still poorly understood. Here we investigate the evolutionary history of an ancient and widespread arachnid lineage (Pseudoscorpiones: Chthoniidae: Tyrannochthoniini) in the WG through a variety of biogeographical and diversification analyses, including sequence data from three markers for 45 terminals from the WG and 22 from Africa, East and Southeast Asia, Australasia and the Neotropics. Our results show that WG Tyrannochthoniini are rendered paraphyletic by African taxa, a result consistent with continental drift. WG Tyrannochthoniini are further split into two major clades, a southern WG clade and an African-central WG clade, due to vicariance along a major biogeographical barrier, the Palghat Gap. Central WG pseudoscorpions diverged from their African relatives at the Jurassic-Cretaceous boundary, coinciding with the separation of India from Africa. Despite ancient vicariance, six dispersal events occurred across the Palghat Gap, beginning in the Early Cretaceous as India was rafting towards Southeast Asia and ending by the early Miocene, implying that the Palghat Gap became impermeable to dispersal for these humid-adapted organisms during Miocene aridification. Although the ancestor of WG Tyrannochthoniini originated in lowland WG forests, three dispersal events into high montane Shola forests occurred most likely as a result of Late Cretaceous orogenesis and Neogene uplift, allowing lineages to occupy niches at higher elevations. An exponentially declining diversification rate, typical of older lineages, supports the 'museum' model of diversification for WG Tyrannochthoniini. Our study sheds light on the historical biogeography of relictual soil-arthropods in the WG and emphasises the role of WG forest refugia in preserving ancient invertebrate biodiversity. [ABSTRACT FROM AUTHOR]

Scorpions possess two types of visual organs, the median and lateral eyes. Both eyes consist of simple ocelli with biconvex lenses that differ in structure and function. There is little variation in the number of median ocelli across the order. Except for a few troglomorphic species in which the median ocelli are absent, all scorpions possess a single pair. In contrast, the number of pairs of lateral ocelli varies from zero to five across Scorpiones and may vary within species. No attempt has been made to homologize lateral ocelli across the order, and their utility in scorpion systematics has been questioned, due to the variation in number. A recent study examined the number of lateral ocelli among various Asian Buthidae C.L. Koch, 1837 and proposed a “five-eye model” for the family. This model has not been examined more broadly within Buthidae, however, nor compared with the patterns of variation observed among other scorpion families. An eyespot, referred to as an accessory lateral eye, situated ventral or posteroventral to the lateral ocelli, has also been reported in some scorpions. Analysis of its structure suggests it serves a nonvisual function. We present the first comparative study of variation in the lateral ocelli across the order Scorpiones, based on examination of a broad range of exemplar species, representing all families, 160 genera (78%), 196 species (9%), and up to 12 individuals per species. We propose a six-ocellus model for Recent scorpions with four accessory ocelli observed in various taxa, homologize the individual ocelli, and correct erroneous counts in the recent literature. We also investigate the presence of the eyespot across scorpions and discover that it is more widespread than previously recognized. Future work should investigate the genetic and developmental mechanisms underlying the formation of the lateral ocelli to test the hypotheses proposed here. [ABSTRACT FROM AUTHOR]

[Display omitted] • Limits of species groups and several species within Buthacus are unclear. • Variation in multilocus DNA and morphology were investigated among more than 30 populations of Buthacus in the Levant. • Traditional Buthacus species groups are invalidated. • Two geographically-delimited clades of species are identified within the genus. • Divergence between the African and Asian clades occurred in the Early Miocene. Sand scorpions of the genus Buthacus Birula, 1908 (Buthidae C.L. Koch, 1837) are widespread in the sandy deserts of the Palearctic region, occurring from the Atlantic coast of West Africa across the Sahara, and throughout the Middle East to Central Asia. The limits of Buthacus , its two species groups, and many of its species remain unclear, and in need of revision using modern systematic methods. The study presented here set out to investigate the phylogeny and biogeography of the Buthacus species occurring in the Levant, last studied in 1980. A phylogenetic analysis was performed on 104 terminals, including six species collected from more than thirty localities in Israel and other countries in the region. Three mitochondrial and two nuclear gene loci were sequenced for a total of 2218 aligned base-pairs. Morphological datasets comprising 22 qualitative and 48 quantitative morphological characters were compiled. Molecular and morphological datasets were analyzed separately and simultaneously with Bayesian Inference, Maximum Likelihood, and parsimony. Divergence time and ancestral range estimation analyses were performed, to understand dispersal and diversification. The results support a revised classification of Levantine Buthacus , and invalidate the traditional species groups of Buthacus , instead recovering two geographically-delimited clades, an African clade and an Asian clade, approximately separated by the Jordan Valley (the Jordan Rift Valley or Syro-African Depression), the northernmost part of the Great Rift Valley. The divergence between these clades occurred in the Early Miocene (ca. 19 Ma) in the Levant, coinciding temporally with the existence of two land bridges, which allowed faunal exchange between Africa and Asia. [ABSTRACT FROM AUTHOR]

More than 85 species of cave-obligate (troglobiotic) millipede have been described from North America. Understanding the patterns and processes that determine their distribution in this region is an area of recent research. Here, we present the first molecular phylogeographic study of troglobiotic millipedes. Millipedes of the genus Tetracion Hoffman, 1956 (Callipodida: Abacionidae) inhabit caves on the Cumberland Plateau in Tennessee and Alabama, a global hotspot for cave biodiversity. Three species have been described: T. jonesi Hoffman, 1956, T. antraeum Hoffman, 1956, and T. tennesseensis Causey, 1959. To examine genetic divergence within and between species of Tetracion we sequenced part of the mitochondrial cytochrome oxidase 1 gene from 53 individuals from eleven caves across the range of T. tennesseensis and in the northern part of the range of T. jonesi. We found: (1) little variation within species (six haplotypes in T. tennesseensis and four haplotypes in T. jonesi, with a maximum of 1.4% intraspecific divergence between haplotypes), (2) that gene flow between caves is limited (7 of 10 haplotypes were restricted to a single cave, and FST > 0.80 and P < 0.05 for fifteen of eighteen comparisons between caves), and (3) significant genetic divergence between species (8.8% between T. tennesseensis and T. jonesi). Our results are consistent with previous morphology-based species definitions showing T. tennesseensis and T. jonesi belonging to distinct taxa. Our research contributes to the growing body of phylogeographic information about cave species on the Cumberland Plateau, and provides a point of comparison for future studies of troglobionts and millipedes. [ABSTRACT FROM AUTHOR]

Aim: Understanding the historical biogeography of the Earth's oldest terrestrial lineages provides insights into lineage diversification in relation to plate tectonics, climate change and biome shifts at maximum timescales. We investigate the biogeography of an ancient arachnid family, dragon pseudoscorpions, which are found today in mesic (mostly temperate) forests on all continents except Antarctica and Europe, have potential origins on Pangea and comprise species with extremely limited dispersal capacities. We evaluate the respective role of continental vicariance (abiotic) and biome shifts (biotic) deep in time and unravel the evolutionary history of this ancient group. Location: Global. Taxon: Pseudotyrannochthoniidae (Arachnida: Pseudoscorpiones). Methods: Five loci were sequenced for 75 Pseudotyrannochthoniidae samples collected across the globe. A matrix was compiled comprising 106 terminals and ~8800 bp, and phylogenetic analyses were performed to uncover relationships. Divergence time and ancestral range estimation analyses were used to reconstruct historical biogeography. Results: Pseudotyrannochthoniidae was monophyletic with high support but relationships among genera did not reflect current taxonomy and instead showed geographical structuring. Pseudotyrannochthoniidae originated in East Asia during the Middle Triassic and began diversifying in the Early Cretaceous. Main Conclusions: Diversity and distributional patterns of dragon pseudoscorpions can be explained by the interplay of continental vicariance through Pangaean breakup, and biome shifts via the spread of temperate habitats in the Cretaceous and their ongoing distributional fluctuations. Pseudotyrannochthoniidae diversification began in the Early Cretaceous as they dispersed with temperate forests across the Northern Hemisphere. Cretaceous vicariance, followed by widespread Holarctic extinction in the Late Tertiary–Quaternary, created their disjunct distribution at northern latitudes. Northern and Southern Hemisphere lineages diverged as Gondwana and Laurasia rifted, and Gondwanan breakup resulted in Afrotropical/Indomalayan and Austral clades. Austral lineages spread with temperate forests, however, cooling caused their disappearance from Antarctica and led to disjunct distributions at low latitudes. [ABSTRACT FROM AUTHOR]

The first integrative systematic revision of the relictual Asian scorpion family Pseudochactidae Gromov, 1998, making use of an unprecedented collection of material acquired during several expeditions to most of the type localities, is presented. The subfamilies, genera and species of Pseudochactidae are revised based on a phylogenetic analysis of 140 morphological characters and 8608 nucleotide base pairs of concatenated DNA sequence from two nuclear and three mitochondrial gene loci, and a multivariate statistical analysis of 22 ratios and 8 counts for 60 specimens. Three subfamilies, four genera and six species are recognized in the family. Troglokhammouaninae, subfam. nov., is created to restore the monophyly of the nominotypical subfamily Pseudochactinae Gromov, 1998. Aemngvantom, gen. nov., is created to accommodate Aemngvantom lao (Lourenço, 2012), comb. nov., and Aemngvantom thamnongpaseuam gen. et sp. nov. Four new synonyms are presented: Troglokhammouanus louisanneorum Lourenço, 2017 = Troglokhammouanus steineri Lourenço, 2007, syn. nov.; Vietbocap thienduongensisLourenço and Pham, 2012 = Vietbocap canhiLourenço and Pham, 2010, syn. nov.; Vietbocap aurantiacusLourenço et al., 2018 = V. canhi, syn. nov.; Vietbocap quinquemiliaLourenço et al., 2018 = V. canhi, syn. nov. Revised diagnoses of the subfamilies, genera and species, with comparative images, a key and distribution maps are provided, along with a summary of available data on ecology and conservation status, where applicable. Among the Southeast Asian pseudochactids, all of which appear to be obligately cavernicolous, the three species of Vietbocapinae Lourenço, 2012, are highly troglomorphic whereas the sole species of Troglokhammouaninae is barely so. Applying recently revised definitions of the Schiner-Racovitza system for the classification of subterranean organisms, only Vietbocapinae can be considered troglobitic. The global diversity of cavernicolous, troglomorphic and troglobitic scorpions is similarly revisited and a key to ecological classification of cavernicolous and troglomorphic scorpions presented. The world totals of troglomorphic vs. troglobitic scorpions are currently 58 vs. 28 species, in 29 vs. 17 genera and 15 vs. 13 families, respectively. [ABSTRACT FROM AUTHOR]

The Malagasy giant pill-millipede genus Sphaeromimus de Saussure & Zehntner, 1902 is revised. Seven new species, S. titanus sp. n., S. vatovavy sp. n., S. lavasoa sp. n., S. andohahela sp. n., S. ivohibe sp. n., S. saintelucei sp. n., and S. andrahomana sp. n. were discovered, in one case with the help of sequence data, in the rainforests of southeastern Madagascar. The species are described using light- and scanning electron microscopy. A key to all 10 species of the genus is presented. All but one (S. andohahela) of the newly discovered species are microendemics each occurring in isolated forest fragments. The mitochondrial COI barcoding gene was amplified and sequenced for 18 Sphaeromimus specimens, and a dataset containing COI sequences of 28 specimens representing all Sphaeromimus species (except S. vatovavy) was analyzed. All species are genetically monophyletic. Interspecific uncorrected genetic distances were moderate (4-10%) to high (18-25%), whereas intraspecific variation is low (0-3.5%). Sequence data allowed the correct identification of three colour morphs of S. musicus, as well as the identity of a cave specimen, which although aberrant in its morphology and colouration, was genetically identical to the holotype of S. andrahoma. [ABSTRACT FROM AUTHOR]

Tetracion jonesi Hoffman, 1956 is a troglomorphic cave-limited abacionid milliped found in caves in the southern Cumberland Plateau of Alabama and Tennessee, USA. Specimens collected in a cave in Franklin Co., Tennessee, were live-extracted with methanol. Gas chromatography/mass spectroscopy of the extract revealed p-cresol as the sole component of the milliped's chemical defense. Three other callipodidan species have been studied and also produce p-cresol. [ABSTRACT FROM AUTHOR]