Outsourcing a visual neuropil - The central visual system of the median eyes of Galeodes granti Pocock, 1903 (Arachnida: Solifugae).

Only a few studies have examined the central visual system of Solifugae until now. To get new insights suitable for phylogenetic analysis we studied the R-cell (or retinula cell) projections and visual neuropils of Galeodes granti using various methods. G. granti possesses large median eyes and rudimentary lateral eyes. In this study, only the R-cells and neuropils of the median eyes were successfully stained. The R-cells terminate in two distinct visual neuropils. The first neuropil is located externally to the protocerebrum directly below the retina, the second neuropil lies in the cell body rind of the protocerebrum, and immediately adjacent is the arcuate body. This layout of the median eye visual system differs from Arachnopulmonata (Scorpiones + Tetrapulmonata). However, there are several similarities with Opiliones. In both, (1) the R-cells are connected to a first and second visual neuropil and not to any other region of the brain, (2) the first neuropil is not embedded in the cell body rind of the protocerebrum, it is rather external to the protocerebrum, (3) the second visual neuropil is embedded in the cell body rind, and (4) the second neuropil abuts the arcuate body. These findings may provide important new characters for the discussion on arachnid phylogeny.

Adaptations for matrotrophy in the female reproductive system in the pseudoscorpion Chelifer cancroides (Chelicerata: Pseudoscorpiones, Cheliferidae).

Pseudoscorpiones (pseudoscorpions, false scorpions) is an order of small terrestrial chelicerates. While most chelicerates are lecithotrophic, that is, embryos develop due to nutrients (mostly yolk) deposited in the oocyte cytoplasm, pseudoscorpions are matrotrophic, that is, embryos are nourished by the female. Pseudoscorpion oocytes contain only a small amount of yolk. The embryos develop within a brood sac carried on the abdominal site of the female and absorb nutrients by a pumping organ. It is believed that in pseudoscorpions nutrients for developing embryos are produced in the ovary during a postovulatory (secretory) phase of the ovarian cycle. The goal of our study was to analyze the structure of the female reproductive system during the secretory phase in the pseudoscorpion Chelifer cancroides, a representative of the family Cheliferidae, considered to be one of the most advanced pseudoscorpion taxa. We use diverse microscopic techniques to document that the nutritive fluid is produced not only in the ovaries but also by the epithelial cells in the oviducts. The secretory active epithelial cells are hypertrophic and polyploid and release their content by fragmentation of apical parts. Our observations also indicate that fertilization occurs in the oviducts. Moreover, in contrast to previous findings, we show that secretion of the nutritive material starts when the fertilized oocytes reach the brood sac and thus precedes formation of the pumping organ. Summing up, we show that C. cancroides exhibits traits of advanced adaptations for matrotrophy due to coordinated secretion of the nutritive fluid by the ovarian and oviductal epithelial cells, which substantially increases the efficiency of nutritive fluid formation. Since the secretion of nutrients starts before formation of the pumping organ, we suggest that the embryos are able to absorb the nutritive fluid also in the early embryonic stages.

Interspecific, ontogenetic, and sexual variation in ozopore morphology among cosmetid harvestmen (Arachnida, Opiliones, Laniatores).

The ozopores of cosmetid harvestmen rest upon lateral projections of the carapace, have simple or highly reduced channels, and are partially obscured by enlarged dorsal processes associated with coxae I and II. Rather than use scent gland secretions to form a chemical shield on the dorsum, the cosmetid harvestman exhibits a unique defensive behavior known as "leg dabbing" in which the distal tip of tarsus I or II is dipped into fluid that accumulate at the base of coxa II and the droplet on the tarsus is pointed toward the predator. Relatively little is known about interspecific variation in ozopore morphology among cosmetid harvestmen. In this study, we used scanning electron microscopy to examine the ozopores of males and females of nine species as well as those of antepenultimate nymphs for two species. Among adults, we found differences between species in the shapes of the ozopores (round or subtriangular), the morphology of the dorsal and lateral channels (if present), and the relative size, shape and armature of the dorsal posterior process (dpp) of coxa I and the dorsal anterior process (dap) of coxa II. Our observations suggest that the morphology of dpp I and dap II could be sources for systematic characters in future phylogenetic studies of the Cosmetidae. We observed ontogenetic differences but relatively little intersexual variation in the morphology of the ozopore. The ozopores of nymphs are generally more oval than those of adults and the opening of the ozopore of the nymph is less obstructed, if at all, by the dorsal coxal processes of legs I-II. These morphological differences suggest that nymphs may use scent gland secretions in a manner different from that of adults.

The sensory equipment of a sandokanid: An extreme case of tarsal reduction in harvestmen (Arachnida, Opiliones, Laniatores).

The study of sensory structures has the potential to provide insights into the natural history and evolution of animals. The sensory structures of arachnids are usually concentrated on the pedipalps (the tritocerebral appendages) or on the distal podomere (tarsus) of the anterior walking legs, the latter being the case for armored harvestmen (Opiliones, Laniatores). Therefore, modifications of the tarsus could have direct impacts on the sensory equipment of these animals. Using scanning electron microscopy, we investigated the sensory equipment in an extreme case of reduction in tarsal articles in the harvestman Sandokan truncatus (Sandokanidae), which bears a single tarsomere in all legs, and the potential consequences of this reduction. Additionally, we review the literature on the natural history of the family Sandokanidae. Tarsomeres of all legs are equipped with gustatory sensilla, mechanoreceptors, and a pore organ, but wall-pored olfactory sensilla are restricted to tarsi I and II. Tarsi II present a higher density of olfactory sensilla and also putative campaniform sensilla (strain detectors), which indicates a special sensory function of this pair of legs. Other podomeres are covered with shelled sensilla, a probable chemoreceptor previously unreported in Opiliones. Overall, S. truncatus has types of sensilla largely comparable to harvestmen with longer and subdivided tarsi. However, S. truncatus also exhibits extra-tarsal sensory fields of sensilla basiconica (putative thermo-/hygroreceptors) in previously undescribed sites, and the unique pore organs. Our results establish a basis for further research investigating the natural history, as well as the evolutionary correlations and mechanistic causes of the tarsal reduction in this enigmatic lineage.

Ontogenetic variation in the sensory structures on the pedipalps of cosmetid harvestmen (Arachnida, Opiliones, Laniatores).

In arachnids, pedipalps are highly variable appendages that may be used in feeding, courtship, defense, and agonistic encounters. In cosmetid harvestmen, adults have pedipalps that feature flattened femora, spoon-shaped tibiae, and robust tarsal claws. In contrast, the pedipalps of nymphs are elongate with cylindrical podomeres and are adorned with delicate pretarsi. In this study, we used scanning electron microscopy to examine the distribution of cuticular structures (e.g., sensilla chaetica, pores) on the elements of the pedipalps of adults and nymphs of three species of cosmetid harvestmen. Our results indicate that there is considerable ontogenetic variation in the morphology of the trochanter, femur, patella, tibia, and tarsus. The pretarsus of the nymph has a ventral patch of setae that is absent from the adult tarsal claw. We observed this structure on all three cosmetid species as well as on the pedipalps of an additional seven morphospecies of nymphs collected in Belize and Costa Rica. This structure may represent a previously unrecognized autapomorphy for Cosmetidae. Examinations of the pedipalps of antepenultimate nymphs of additional gonyleptoidean harvestmen representing the families Ampycidae, Cranaidae, Manaosbiidae, and Stygnidae revealed the occurrence of unusual, plumose tarsal setae, but no setal patches on the tarsal claw.

Ultrastructure of chemoreceptive tarsal sensilla in an armored harvestman and evidence of olfaction across Laniatores (Arachnida, Opiliones).

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.

The integumentary ultrastructure of Cryptocellus bordoni Dumitresco and Jurvara-Bals, 1976 (Arachnida, Ricinulei).

Ricinulei is an order of Arachnida composed of rare and little known species. The species of Ricinulei possess a rich variety of fine integumentary structures that have been poorly investigated in a few species. Besides, several structures are still undescribed and their function not yet addressed. In this paper we provide a detailed study of the integumentary morphology of Cryptocellus bordoni Dumitresco and Jurvara-Bals, 1976 using Light and Scanning Electron Microscopy. We describe and present photos of the new and already known fine integumentary structures. We compare the new structures to those previously described for other Ricinulei species and discuss their taxonomic implications and the placement of C. bordoni in the genus Cryptocellus.

Functional anatomy of the pretarsus in whip spiders (Arachnida, Amblypygi).

Whip spiders (Amblypygi) are a small, cryptic order of arachnids mainly distributed in the tropics. Some basal lineages (families Charinidae and Charontidae) have adhesive pads on the tips of their six walking legs. The present study describes the macro- and ultrastructure of these pads and investigates their contact mechanics and adhesive strength on smooth and rough substrates. Furthermore, the structure of the pretarsus and its kinematics are compared in Charon cf. grayi (with an adhesive pad) and Phrynus longipes (without an adhesive pad). The adhesive pads exhibit an elaborate structure with a unique combination of structural features of smooth and hairy foot pads including a long transversal contact zone performing lateral detachment, a thick internally-branched cuticle with longitudinal ribs and hexagonal surface microstructures with spatulate keels. The contact area of the pads on smooth glass is discontinuous due to the spatulate microstructures with a discontinuous detachment, which could be observed in vivo by high speed videography at a rate of up to 10,000 fps. Adhesive strength was measured with vertical whole animal pull-off tests, obtaining mean values between 55 and 200 kPa. The occurrence of viscous lipid secretions between microstructures was occasionally observed, which, however, seems not to be a necessity for good foothold. The results are discussed in relation to the whip spider's ecology and evolution. Structure-function relationships of the adhesive pads are compared to those of insects and vertebrates.

A new epigean harvestman of the genus Guerrobunus (Laniatores: Phalangodidae), from Hidalgo, Mexico, with taxonomic notes about the genus.

The fifth species of the genus Guerrobunus, Guerrobunus barralesi sp. nov. is described from specimens collected in Hidalgo, Mexico. This species represents the first eastern record from the country and unlike other species of the genus, is completely epigean. Guerrobunus barralesi sp. nov. is compared with the most similar species, Guerrobunus minutus, which also has well developed and pigmented eyes. Finally, external morphology, including male genitalia, taxonomy of the genus, and familial assignment into the family Phalangodidae are discussed.

A taxonomic revision of the Neotropical spider genus Xiruana Brescovit 1997 (Araneae: Anyphaenidae, Anyphaeninae).

The genus Xiruana Brescovit, 1997 is currently composed of four South American species: X. gracilipes (Keyserling) from Brazil, Bolivia and Argentina, X. affinis (Mello-Leitão) from Brazil, X. hirsuta (Mello-Leitão) from Venezuela, Brazil, Paraguay, Argentina and Uruguay, and X. tetraseta (Mello-Leitão) from Venezuela, Brazil and Paraguay. Of these, the last three are redescribed in this paper, including the first description of the females of X. hirsuta and X. tetraseta. Additionally, we describe thirteen new species: Xiruana pocone n. sp. from Brazil, Paraguay and Argentina; X. bifida n. sp. from Brazil and Paraguay; X. aymara n. sp. from Bolivia; X. cocha n. sp. from Peru; X. fiebrigi n. sp. from Paraguay, and X. ajuricaba n. sp., X. tribarrense n. sp., X. guaia n. sp., X. jaboticabal n. sp., X. minacu n. sp., X. tapirape n. sp., X. lusitania n. sp., X. silarae n. sp., all endemic to Brazil. The known geographical distribution of all species here presented is mapped.

How to stay on mummy's back: Morphological and functional changes of the pretarsus in arachnid postembryonic stages.

A specific type of maternal care occurs in several groups of Arachnida: mothers carry their offspring on their back (pulli-carrying behaviour). In scorpions, whip scorpions and whip spiders it is the prenymphal stage that settles on the mother. The prenymph is not yet fully developed for a free life and very limited in its mobility, but its feet are equipped with special adhesive organs (arolia) that become lost at the nymphal stage. Here we study the morphology, ultrastructure and mechanical function of the arolia. In scorpions (Scorpiones) the contact area between arolia and substrate and thus adhesion of the pad is controlled by the antagonistic work of hydrostatic pressure and muscular retraction. Arolia of whip scorpions (Thelyphonida) do not require muscular action for strong attachment. Arrays of long, branching fibres in the mesocuticle lead to high compliancy of the pad. In whip spiders (Amblypygi) the prenymphal pretarsus is already equipped with sclerites and claws. Its arolium is retained in nymphs and adults in some taxa, but acquires a more complex structure. These results contribute to our knowledge on the postembryonic development of arachnids and to the understanding of attachment pad evolution among arthropods. Some of the described developmental, structural, and mechanical phenomena are not known from other animals and might be of potential interest for further biomimetic developments.

Changes in the midgut diverticula in the harvestmen Amilenus aurantiacus (Phalangiidae, Opiliones) during winter diapause.

The harvestmen Amilenus aurantiacus overwinter in diapause in hypogean habitats. The midgut diverticula have been studied microscopically (light microscopy, TEM) and biochemically (energy-storing compounds: lipids and glycogen) to analyze changes during this programmed starvation period. Throughout the investigated period, the epithelium of the midgut diverticula is composed of secretory cells, digestive cells and adipocytes. Additionally, after the middle of overwintering, the excretory cells appear, and two assemblages of secretory cells are present: the SC1 secretory cells are characterized by electron-dense cytoplasm with numerous protein granules, and the SC2 cells by an electron-lucent cytoplasm with fewer protein granules. The autophagic activity is observed from the middle of overwintering, indicating its vital role in providing nutrients during this non-feeding period. Lipids and glycogen are present in the midgut diverticula cells, except in the excretory cells. Measurements of the lipid droplet diameters and the lipid quantities yielded quite comparable information on their consumption. Lipids are gradually spent in both sexes, more rapidly in females, owing to ripening of the ovaries. Glycogen rates decrease towards the middle, and increase just before the end of overwintering, indicating that individuals are preparing for the epigean active ecophase.

Interspecific variation in the microanatomy of cosmetid harvestmen (Arachnida, Opiliones, Laniatores).

Scanning electron microscopy (SEM) is a useful tool for identifying interspecific variation in often overlooked structures that may represent useful sources for informative phylogenetic characters. In this study, we used SEM to compare the morphology of 12 cosmetid species from Central America, the Caribbean, and North America including multiple species for the genera Cynorta, Erginulus, and Paecilaema. To determine if microanatomical structures were unique to the cosmetid taxa under examination, we investigated the microanatomical structures of six additional species of gonyleptoidean harvestmen representing the families Agoristenidae, Cranaidae, Gonyleptidae, Manaosbiidae, and Stygnidae. Our results indicate that the shape of the ocularium (narrow, intermediate, or broad) did not vary within cosmetid genera, whereas the morphology of the rough pit glands on the eye mound varied considerably between species. Each cosmetid species had 10-20 rough pit glands on the ocularium whereas only the eye mounds of Avima intermedia (Agoristenidae) and Glysterus sp. (Gonyleptidae) had similar structures. With regards to the surface texture of the dorsal scutum, cosmetid harvestmen exhibited a rivulose-microgranulate morphology (6 species), a microtuberculate-rivulose-microrgranulate morphology (4 species), or a microgranulate morphology (2 species). In contrast, each of the gonyleptoidean species exhibited a microgranulate pattern, with the exception of Stygnoplus clavotibialis, which had a rivulose-microgranulate surface texture. For cosmetid harvestmen, we observed considerable interspecific variation in the shape and number of teeth on the fixed and moveable fingers of the male chelicerae. Similarly, we also observed interspecific variation in the distribution and shape of tubercles on the ventral and dorsal surfaces of the femur of the pedipalp. Overall, our results indicate that there are several microanatomical structures associated with the ocularium, dorsal scutum, male chelicera, and pedipalp that could represent informative phylogenetic characters in future taxonomic studies of cosmetid harvestmen.

Ovipositor morphology of cosmetid harvestmen (Arachnida, Opiliones, Laniatores): a new source of informative characters.

The external morphology of the penis is an important source of systematic characters in phylogenetic studies of harvestmen. Modern taxonomic studies generally include micrographs generated by scanning electron microscopy (SEM) to illustrate penis morphology. In contrast, the external morphology of the ovipositor has largely been ignored for harvestmen belonging to the suborder Laniatores. Comparative studies of ovipositor microanatomy using SEM are especially lacking for species belonging to the superfamily Gonyleptoidea. In an effort to determine if the ovipositor could be a useful source of informative characters for these harvestmen, we investigated interspecific variation in the external morphology of the ovipositor for 14 species from the family Cosmetidae. Our SEM-based study revealed that the external surface of the distal tips of the ovipositors of most species was generally divided into four symmetrical lobes, although we observed a bilobed condition in Erginulus clavotibialis and Erginulus subserialis. The distal surfaces were also generally smooth, with the exception of the ovipositor of Erginulus weyerensis, which featured small surface setae. In addition, we observed considerable interspecific variation in the morphology of the peripheral setae on the distal tip, especially with respect to relative size, morphology of the shaft, and number, symmetry, and shapes of the distal tips. The functional significance, if any, of variation in the structure of the peripheral setae is unclear. Additional behavioral studies of copulation and oviposition are needed to determine the functional relationships between reproductive morphology and behavior. The morphological variation that we observed suggests that future taxonomic studies of cosmetid harvestmen, and potentially other gonyleptoidean taxa, would benefit from the inclusion of descriptions of ovipositor morphology.

Autophagic activity in the midgut gland of the overwintering harvestmen Gyas annulatus (Phalangiidae, Opiliones).

Juvenile harvestmen Gyas annulatus overwinter in dormancy in hypogean habitats for 4-5 months. The ultrastructure of the autophagic structures in their midgut epithelium cells was studied by light microscopy, transmission electron microscopy (TEM) and immunofluorescence microscopy (IFM) during this non-feeding period. Before overwintering (November), autophagic structures were scarce. In the middle (January) and at the end of overwintering (March), phagophores, autophagosomes and autolysosomes were present in the cytoplasm of both the secretory and the digestive midgut epithelium cells, gradually increasing their abundance during overwintering. In addition, vacuolization of the cytoplasm intensified. Both processes are induced by starvation. Autophagic structures and cytoplasm vacuolization enable the reuse of the cell's own components required for the maintenance of vital processes during dormancy. While TEM is a much more convenient method for recognition of the autophagic structure types and their ultrastructure, IFM enables exact counting of these structures.

Ultrastructure of the sexually dimorphic tarsal glands and tegumental glands in gonyleptoid harvestmen (Opiliones, Laniatores).

In at least four closely related families of the diverse harvestmen lineage Gonyleptoidea, males may possess sexually dimorphic tarsal glands in the swollen tarsomeres of the basitarsus and/or metatarsus of leg I. The first histological and ultrastructural examination of the sexually dimorphic tarsal glands in leg I focused only on Manaosbiidae. In this study, we examine the morphology and ultrastructure of the sexually dimorphic glands, and their associated glandular openings, found in the basitarsus and/or metatarsus of leg I of males representing Cosmetidae, Gonyleptidae, and Cranaidae (glandular openings only). In cosmetids and gonyleptids, the tarsal glands are made up of 20­60 glandular units that form distinct groups within the prolateral and retrolateral half of the tarsomere. Each glandular unit consists of a pair of terminal secretory cells, an intercalary cell wrapped around the receiving canal, and a canal cell tightly wrapped around the length of the conducting canal. Cosmetidae, Gonyleptidae, and Cranaidae exhibit remarkably similar tarsal glands and gland openings although the location of the glands in the leg differs slightly among them. Males of these three families exhibit markedly different glands and glandular openings compared to males of the family Manaosbiidae. The sexually dimorphic tarsal glands may provide an important morphological character for determining phylogenetic relationships among gonyleptoid families. Finally, we provide morphological and ultrastructural data for the common tegumental glands. These data indicate that the sexually dimorphic tarsal glands are strikingly similar to, and may possibly be derived from, the tegumental glands.

The harvestman tarsus and tarsal flexor system with notes on appendicular sensory structures in Laniatores.

The tarsal flexor system, a novel system of retinacular structures, is described for the first time based on morphological and ultrastructural examinations of several Neotropical harvestmen (Opiliones: Laniatores). The tarsal flexor system is made up of many individual pulleys that function to maintain close apposition between the tendon and internal ventral surface of the cuticle in the tarsus. Pulley cells are specialized tendinous cells that form the semi-circular, retinacular pulley system in the tarsus; these cells contain parallel arrays of microtubules that attach to cuticular fibers extending from deep within the cuticle (i.e., tonofibrillae). The tarsal flexor system is hypothesized to provide mechanical advantage for tarsal flexion and other movements of the tarsus. This system is discussed with regards to other lineages of Opiliones, especially those that exhibit prehensility of the tarsus (i.e., Eupnoi). Comparing tarsal morphology of laniatorid harvestmen to other well-studied arachnids, we review some literature that may indicate the presence of similar tarsal structures in several arachnid orders. The general internal organization of the tarsus is described, and ultrastructural data are presented for a number of tarsal structures, including sensilla chaetica and the tarsal perforated organ. Sensilla chaetica possess an internal lumen with dendritic processes in the center and exhibit micropores at the distal tip. With respect to the tarsal perforated organ, we found no ultrastructural evidence for a sensory or secretory function, and we argue that this structure is the result of a large pulley attachment site on the internal surface of the cuticle. A small, previously undocumented muscle located in the basitarsus is also reported.

Description of the male, larva and nymphal stages of Cryptocellus iaci (Arachnida, Ricinulei), with an overview of tarsal sensilla and other integumental structures.

The male, larva and nymphal stages of Cryptocellus iaci Tourinho, Lo Man-Hung & Bonaldo, 2010, a species previously known only from a single female, are described based on specimens from around the type locality, in an area of both Terra Firme forest and igapó (flooded forests), at the Jufari River, Roraima State, Brazil. The specimens were illustrated using live photography, stereomicroscopy, and scanning electron microscopy, allowing us to examine and describe the large diversity of tarsal sensilla and other integumental structures in Cryptocellus and to compare them to those of the previously studied Pseudocellus. Based on the male somatic characters Cryptocellus iaci is placed in thefoedus species-group. Cryptocellus iaci has two sensilla of type 1 on the distal tarsomeres of legs III (DT III), while only one has been reported for Pseudocellus spp., suggesting a potential value in this type of character for systematic studies of the group.

Two new species of ricinuleids of the genus Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from southern Mexico.

Two new species of ricinuleids of the genus Pseudocellus are described from Mexico: Pseudocellus cruzlopezi sp. nov. from Oaxaca, and Pseudocellus monjarazi sp. nov. from Chiapas. Both species are described from adult males and females. The first species is epigean and edaphomorphic, whereas the second is cavernicolous and troglomorphic. The number of known species of the genus Pseudocellus increases to 25, and Mexican species to 16, indicating that Mexico has the highest diversity of ricinuleids in the world. An updated identification key to adult males of the 16 described species found in Mexico and southern USA is provided.

Novel discovery of lamellar papillae on the grooming organ in Synsphyronus (Garypidae: Pseudoscorpiones).

The chelicerae, the first pair of appendages in Chelicerata, exhibit morphological and functional variation across arachnid orders. The two-segmented chelicerae of pseudoscorpions serve multiple functions including grooming, courtship, and grasping prey. Scanning electron microscopy was used to investigate structures found on the chelicera; the serrulae interiores and exteriores, grooming organs, were given special attention. Functional analogies were made between the cheliceral structures documented in pseudoscorpions and those found in other arthropods. The novel discovery of vestitural papillae and a patch of elongate papillae on the serrula exterior are reported. The focal taxon for the study is Synsphyronus (Garypidae), an Australasian genus.