Search

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.

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]

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]

[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]