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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.

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.

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.

Brill profit levels and organic growth in line with expectations, integration V&R according to plan and continuation of eBusiness growth Update on year-to-date performance During the third quarter Brill's eBusiness continued its strong growth, driven by eBooks and digital primary sources. This growth compensated for the ongoing decline in print books. YTD Journal sales declined mainly due to delayed invoicing, which is expected to be recovered in Q4, and continuing IT problems at our UK based distributor. Overall Brill's revenues including the acquisition of Vandenhoeck & Ruprecht and Böhlau Verlag (V&R) are clearly ahead of last year.Cost of goods sold improved slightly as a result of the migration from print to eBooks. Operating costs were in line with expectations. Year to date EBITDA, operating profit and net profit have developed as planned.The integration of V&R runs according to plan. Both the Brill as well as the V&R teams and staff are now part of one global matrix organization. Various integration projects in IT, production, distribution, publishing and sales are underway. Management closely leads and monitors the integration and is pleased with the progress and the results of the acquired V&R business so far.In August, Brill acquired the journal Folia Primatologica from Karger Publishing, an important addition to our Biology portfolio. At the end of Q3 Brill's Book Archive was launched, a digital archive expanding our eBook list with more than 6,000 back list titles in 2021. The first sales are expected in Q4. FY OutlookThe COVID-19 pandemic continues to make market circumstances difficult and unpredictable. In Asia important countries are still in lockdown and (intercontinental) travel is still difficult or impossible, limiting our staff to visit authors, conferences and customers in person. The shortages in the global supply chains are hampering Brill as well, to the extent that this may have a negative impact on the results of the important 4 th quarter and could affect our top-and bottom line negatively.

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CONTENTS SIAC gih.Sh o: Petar scat NG gael Marit ts aad oe tle s.r aeet sca de Shale antl Maid te gud ce Vath teal 3 1ACURMOT NU VGLa1S) eR org OR i eT ARRON cede lee arene ated ee hut RE a da ed eee MAME dee 3 Review of Cavernicolous, Troglobitic, and Troglomorphic Scorpions .