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Spiders are among the most diverse animals, which developed different morphological and behavioral traits for capturing prey. We studied the anatomy and functionality of the rare and apomorphic raptorial spider feet using 3D reconstruction modeling, among other imaging techniques. The evolutionary reconstruction of the raptorial feet (tarsus plus pretarsus) features using a composite tree of spiders, indicating that similar traits emerged three times independently in Trogloraptoridae, Gradungulinae, and Doryonychus raptor (Tetragnathidae). The characteristics defining the raptorial feet are an interlocked complex merging of the base of the elongated prolateral claw with the pretarsal sclerotized ring, with the former clasping against the tarsus. Raptorial feet even flex over robust raptorial macrosetae forming a reduced tarsal version of a catching basket to encase prey during hunting. Our results show that Celaeniini (Araneidae) and Heterogriffus berlandi (Thomisidae), taxa previously compared with raptorial spiders, lack the raptorial feet key characteristics and the tarsal-catching basket. We make predictions about the possible behavior of the abovementioned taxa that will need to be tested by observing living specimens. We conclude that multiple morphological tarsal and pretarsal micro-structures define the raptorial foot functional unit and recommend a comprehensive evaluation before assigning this configuration to any spider taxa., Competing Interests: Declaration of competing interest None., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Harvestmen (Arachnida, Opiliones) are especially dependent on chemical cues and are often regarded as animals that rely mainly on contact chemoreception. Information on harvestman sensilla is scarce when compared to other arachnid orders, especially concerning internal morphology. Using scanning (SEM) and transmission (TEM) electron microscopy, we investigated tarsal sensilla on the distal tarsomeres (DT) of all leg pairs in Heteromitobates discolor (Laniatores, Gonyleptidae). Furthermore, we explored the typological diversity of sensilla present on the DT I and II in members of the suborder Laniatores, which include two thirds of the formally described opilionid fauna, using species from 17 families representing all main laniatorian lineages. Our data revealed that DT I and II of H. discolor are equipped with wall-pored falciform hairs (two types), wall-pored sensilla chaetica (two types) and tip-pored sensilla chaetica, while DT III and IV are mainly covered with trichomes (non-sensory) and tip-pored sensilla chaetica. The ultrastructural characteristics support an olfactory function for all wall-pored sensilla and a dual gustatory/mechanoreceptive function for tip-pored sensilla chaetica. Based on our comparative SEM survey, we show that wall-pored sensilla occur in all investigated Laniatores, demonstrating their widespread occurrence in the suborder and highlighting the importance of both legs I and II as the sensory appendages of laniatorean harvestmen. Our results provide the first morphological evidence for olfactory receptors in Laniatores and suggest that olfaction is more important for harvestmen than previously thought., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The superfamily Austrochiloidea (Austrochilidae and Gradungulidae) take a pivotal position in araneomorph spider phylogeny. In this discussion crevice weaver spiders (Filistatidae) are of equal interest. Especially data from these phylogenetically uncertain yet basal off branching groups can enlighten our understanding on the evolution of organ systems. In the course of a survey on the evolutionary morphology of the circulatory system in spiders we therefore investigated the hemolymph vascular system in two austrochiloid and one filistatid species. Additionally some data on a hypochilid and a gradungulid species are included. Using up-to-date morphological methods, the vascular systems in these spiders are visualized three dimensionally. Ground pattern features of the circulatory systems in austrochiloid spiders are presented and the data discussed along recent lines of phylogenetic hypotheses. Special topics highlighted are the intraspecific variability of the origins of some prosomal arteries and the evolutionary correlation of respiratory and circulatory systems in spiders., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The male reproductive system and spermatozoa of spiders are known for their high structural diversity. Spider spermatozoa are flagellate and males transfer them to females in a coiled and encapsulated state using their modified pedipalps. Here, we provide a detailed overview of the present state of knowledge of the primary male reproductive system, sperm morphology and the structural diversity of seminal fluids with a focus on functional and evolutionary implications. Secondly, we conceptualized characters for the male genital system, spermiogenesis and spermatozoa for the first time based on published and new data. In total, we scored 40 characters for 129 species from 56 families representing all main spider clades. We obtained synapomorphies for several taxa including Opisthothelae, Araneomorphae, Dysderoidea, Scytodoidea, Telemidae, Linyphioidea, Mimetidae, Synotaxidae and the Divided Cribellum Clade. Furthermore, we recovered synspermia as a synapomorphy for ecribellate Haplogynae and thus propose Synspermiata as new name for this clade. We hope that these data will not only contribute to future phylogenetic studies but will also stimulate much needed evolutionary studies of reproductive systems in spiders., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Secondary sexual traits in males can extend to glandular structures that play a role during courtship and mating. In dwarf spiders (Linyphiidae, Erigoninae), glandular secondary sexual traits are particularly common. Males are characterized by cephalic modifications which produce secretions that females contact with their mouthparts during courtship and/or copulation. We used the dwarf spider Oedothorax retusus as a model species to investigate if and when the contents of the glands are released during a mating sequence and if so, if the gland reservoirs are refilled after mating. To this aim, we quantitatively compared the glandular tissue on the ultrastructural level between a) inexperienced males, b) males that performed courtship, c) males immediately after copulation, and d) males three days after mating. We assessed whether the treatment groups differed in the filling state of the conducting canals and receiving canals (reservoir regions) of the glandular units. Our study shows that courting males as well as males three days after mating did not differ significantly from control (inexperienced) males in the presence of secretions. However, males exhibited significantly less secretion immediately after mating. This strongly suggests that the main function of the secretions is gustatorial courtship and not the emission of volatile pheromones for mate attraction as was previously assumed., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Sperm conjugation, where two or more sperm are physically united, is a rare but widespread pheno-menon across the animal kingdom. One group well known for its different types of sperm conjugation are spiders. Particularly, haplogyne spiders show a high diversity of sperm traits. Besides individual cleistospermia, primary (synspermia) and secondary (coenospermia, "spermatophore") sperm conjugation occurs. However, the evolution of sperm conjugates and sperm is not understood in this group. Here, we look at how sperm are transferred in Caponiidae (Haplogynae) in pursuit of additional information about the evolution of sperm transfer forms in spiders. Additionally, we investigated the male reproductive system and spermatozoa using light- and transmission electron-microscopy and provide a 3D reconstruction of individual as of well as conjugated spermatozoa. Mature spermatozoa are characterized by an extremely elongated, helical nucleus resulting in the longest spider sperm known to date. At the end of spermiogenesis, synspermia are formed by complete fusion of four spermatids. Thus, synspermia might have evolved early within ecribellate Haplogynae. The fused sperm cells are surrounded by a prominent vesicular area. The function of the vesicular area remains still unknown but might be correlated with the capacitation process inside the female. Further phylogenetic and functional implications of the spermatozoa and sperm conjugation are discussed., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

This study is the first report on the male genital system and the sperm structure of the South American genus Cryptocellus and provides a second description for the Central American genus Pseudocellus. The spermatids of the Colombian species Cryptocellus narino are elongated and anteriorly lentoid-shaped due to two conspicuous intracellular electron-dense plates. Two cell protrusions are present, which contain in front of the lentoid part the acrosomal complex and parts of the axoneme and nucleus, and behind the lentoid part the continuing axoneme and nucleus. The acrosomal filament originates from a cap-like acrosomal vacuole, extends into the nuclear canal and ends behind the lentoid part. The nucleus runs parallel to the axoneme. The axoneme possesses a typical 9+2 microtubular pattern. At the end of spermiogenesis the acrosomal complex, nucleus and axoneme coil within the cell forming cleistospermia as transfer form. Our results of Pseudocellus pearsei confirm an earlier study on that genus which is considered to be not closely related to Cryptocellus. According to the present study the sperm structure of the observed Cryptocellus species is very similar to what is described for Pseudocellus.

The present study deals with the spermatozoa and spermiogenesis of Liphistius cf. phuketensis, a representative of the most primitive and enigmatic spider group Mesothelae. The general organization of the spermatozoa is very similar to the condition known from Amblypygi supporting a sister-group relationship between Araneae and Amblypygi. Besides plesiomorphic characters such as, e.g., an elongated and corkscrew shaped nucleus, the sperm cells are characterized by several apomorphic characters, e.g., the giant body and conspicuous membranous areas which are formed at the end of spermiogenesis. As the transfer form, coenospermia are formed at the end of spermiogenesis, which strongly supports the idea that this type of sperm aggregation is the primitive transfer form within spiders. A very remarkable character of the spermatozoa of some groups of arachnids is the coiling of the main cell organelles at the end of spermiogenesis. Previously, the Mesothelae were believed to be the only spider group which does not show a complete coiling of the main cell organelles. With the present study the first evidence of a complete coiling of spermatozoa within this primitive spider group could be documented, indicating that this character is part of the ground pattern of spider spermatozoa. Consequently, the incomplete coiling seems to be a synapomorphy of certain species of Mesothelae, which sheds new light on the discussion of the phylogenetic relationships of this group.

Two species of the early derivative spider family Heptathelidae (Heptathela kimurai yanbaruensis and Ryuthela nishihirai nishihirai) have been investigated with respect to spermiogenesis, focussing on late events during which peculiar transfer forms are developed. It is shown, for the first time in detail, that these spiders produce coenospermia. The coenospermia of these species are large aggregates containing more than 20 individual encapsulated spermatozoa. The coenospermia possess a likely flexible envelope formed by a thick multilayered secretion, which protects the spermatozoa during transfer to the female genital system and storage in the receptacula. In addition, a short description of the main cell components of the individual spermatozoa is given as a complement to previous studies. With the observation presented here, the suggestion that coenospermia are an ancestral character in the Araneae is further confirmed, and plesiomorphic features of spider sperms are consolidated.