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.     

First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata)

The phylogenetic relationships of the major lineages of the arachnid order Pseudoscorpiones are investigated for the first time using molecular sequence data from two nuclear ribosomal genes and one mitochondrial protein-encoding gene. The data were analyzed using a dynamic homology approach with the new program POY v.4 under parsimony as the optimality criterion. The data show monophyly of Pseudoscorpiones as well as many of its superfamilies (Feaelloidea, Chthonioidea, Cheiridioidea and Sternophoroidea), but not for Neobisiodea or Garypoidea. Cheliferoidea was not monophyletic either due to the position of Neochelanops, which grouped with some garypoids. In all the analyses, Feaelloidea constituted the sister group to all other pseudoscorpions; Chthonioidea is the sister group to the remaining families, which constitute the group Iocheirata sensu Harvey--a clade including pseudoscorpions with venom glands within the pedipalpal fingers. This phylogenetic pattern suggests that venom glands evolved just once within this order of arachnids.     

Anatomy and affinities of a new 535‐million‐year‐old medusozoan from the Kuanchuanpu Formation,South China

We describe here Sinaster petalon gen. et sp. nov., a new embryonic form from the c. 535 million‐year‐old Kuanchuanpu Formation of South China (Ningqiang, Shaanxi Province). The excellent three‐dimensional, phosphatic preservation of these microfossils allowed us to use x‐ray microtomographic techniques to make accurate reconstructions of their internal structures and to compare their anatomy point‐by‐point with that of extant cnidarians and other animal groups. Sinaster petalon has anatomical features typical of extant Medusozoa (Cnidaria), such as coronal muscles, perradial and adradial frenula, interradial septa, accessory septa, gonad‐lamellae, tentacle buds and perradial pockets. Although Sinaster cannot be straightforwardly assigned to any crown‐group within Medusozoa, the presence of marginal lappets and endodermal lamellae suggests that it is closer to Cubozoa and Scyphozoa than to any other group of modern cnidarians. The tentative placement of Sinaster within the stem‐group Cubozoa is justified by the presence of a velarium supported by a frenulum. The cubozoan affinities of Sinaster are also supported by cladistic analysis.     

中国伪蝎名录（蛛形纲：伪蝎目）

本文根据我国伪蝎目已有种类记述,归纳并列出10科34属73种（包括亚种）伪蝎的中文名、拉丁学名、引证及其地理分布,以供今后研究参考。     

LECLERCQIA COMPLEXA: EARLIEST LIGULATE LYCOPOD (MIDDLE DEVONIAN)

Our discovery of a ligule on Leclercqia complexa Banks, Bonamo and Grierson 1972 is the earliest occurrence of a ligulate lycopod in the fossil record. The ligule 1) occurs on a homosporous lycopod, differing with current concepts that the ligule is linked with the heterosporous condition; 2) is located on the leaf far distant from the attachment of leaf to stem, thus differing in position from any known ligulate lycopod, extinct or extant; 3) is comparable in morphology to ligules of extant lycopods, therefore providing no clues as to any earlier specialized function. These findings extend the enigma of the function of the ligule back in time, but necessitate a re-evaluation of the spatial relationship of the leaf and the ligule and of the link between heterospory and the ligule.     

Seasonal aspects of the life cycle of solifuges (Arachnida, Solifugae) as compared with pseudoscorpions (Arachnida, Pseudoscorpiones)

Solifuges (order Solifugae) and pseudoscorpions (order Pseudoscorpiones) united into the superorder Haplocnemata (Shultz, 2007) are nevertheless characterized by essential differences both in morphological and biological characters. Analysis of available information on the biology and life cycles of these arachnids revealed a clear difference between the daily rhythms of activity: their presence in solifuges and their absence in pseudoscorpions. However, this concerning the seasonal adaptations in the two orders is not simple since they demonstrate not only differences but also a lot of similarities. All the studied solifuges are characterized by the seasonally timed stenochronous (heterodynamic) type of development which is characteristic of species with uni-, bi-, and semi-voltine development (i.e., to life cycles completed within a year, half a year, and several years), as well as to species combining different forms of voltinism. This type of development is not only prevalent in solifuges (as in pseudoscorpions and other arachnids) but appears to be the only one, since no cases of eurychronous (homodynamic) development have been found in solifuges; whereas pseudoscorpions and other arachnids develop both steno- and eurychronously. The initial ontogenetic stages remain in underground shelters (brood burrows in solifuges and brood chambers in pseudoscorpions). The first nymphal stages (I instar nymphs in solifuges, protonymphs in pseudoscorpions) are embryonized; active life outside the brood burrows starts with II instar nymphs in solifuges and with deutonymphs in pseudoscorpions.     

Mouthparts in Leptotrombidium larvae (Acariformes: Trombiculidae)

Mouthparts of Leptotrombidium larvae (Acariformes: Trombiculidae), potential vectors of tsutsugamushi disease agents, were studied in detail using light microscopy, scanning electron microscopy, and transmission electron microscopy. The mouthparts incorporated within the pseudotagma gnathosoma are composed of the infracapitulum ventrally and the chelicerae dorsally. The ventral wall of the infracapitulum is formed by a wide mentum posteriorly and a narrowed malapophysis anteriorly. The malapophysis firmly envelops the distal cheliceral portions by its lateral walls. The lateral lips of the malapophysis are flexible structures hiding the cheliceral blades in inactive condition and turning back forming a type of temporary sucker closely applied to the host skin during feeding. The roof of the infracapitulum is formed by a weakly sclerotized labrum anteriorly and a cervix with the capitular apodemes extending posteriorly. The labral muscles are lacking. The capitular apodemes serve as origin for pharyngeal dilators running to the dorsal wall of the pharynx fused with the bottom of the infracapitulum. The basal cheliceral segments are separated from each other besides the very posterior portions where they are movably joined by the inner walls. The sigmoid pieces serve for insertion of the cheliceral elevators originating at the posterior portions of the basal segments. The movable digits reveal the solid basal sclerite and the cheliceral blade curved upward with a tricuspid cap on its tip. Dendrites of nerve cells run along the digits to their tips. The ganglia are placed within the basal segments just behind the movable digits. The chelicerae also reveal well developed flexible fixed digits overhanging the basal portions of the blades. The gnathosoma possesses several sets of extrinsic muscles originating at the scutum and at the soft cuticle behind it. Laterally, the gnathosoma bears five‐segmented palps with a trifurcate palpal claw. J. Morphol. 277:424–444, 2016. © 2016 Wiley Periodicals, Inc.     

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.     

The first cytogenetic characterization of atemnids: pseudoscorpions with the highest chromosome numbers (arachnida: pseudoscorpiones)

The karyotypes of pseudoscorpions of the family Atemnidae (Arachnida: Pseudoscorpiones) were studied for the first time. Karyotype data for 7 species have been obtained. The diploid chromosome numbers of most species considerably exceed the numbers reported in pseudoscorpions so far, with males ranging between 65 and 143. In spite of this, the sex chromosome system of atemnids is characterized by the same features that are found in the majority of other pseudoscorpions with an X0 system; the X chromosome is metacentric and is the largest chromosome or one of the largest chromosomes of the karyotype. Male meiotic cells of Atemnus politus contain 1 or 2 autosome multivalents; most specimens had 2 multivalents. The multivalents were composed of 4, 6, 8 or 10 chromosomes. Multivalent number and structure was consistent within each of the studied individuals. The same number of chromosomes in all of the males examined suggests that multivalents are generated by reciprocal translocations. The high diversity of multivalents suggests considerable range of translocation heterozygosity in the studied population.     

Fine structure of the chelicera in the spider Nephila clavata

The fine structural characteristics of the biting apparatus in the orb‐web spider Nephila clavata were studied using scanning electron microscopy. The main biting apparatuses of spiders are the chelicerae and cheliceral fangs in the cephalothorax. The chelicera of N. clavata is that of the jack‐knife (folding knife) type, which is composed of two segments, and has a labidognathous form that moves at right angles to the body axis. Each chelicera bears a hinged fang that folds into a cheliceral groove. The tips of the fangs are quite sharp, and the spider's body is well adapted to driving the fangs into prey. Just below the fang, each side of the cheliceral groove is covered with a total of seven cuticular teeth (four promarginal teeth and three retromarginal teeth) in two rows. The cheliceral fang has a single aperture at the tip of the posterior surface, and the lower margin of the fang which meets the promarginal teeth is a saw‐like groove. Fine structural observation reveals that each fang has a single venom pore, and each cuticular depressive area on the cheliceral groove has two different types of surface pit. Approximately 40 to 50 spiky protrusions were counted at the cheliceral groove, to hold prey tightly.     

A new microinvertebrate with features of mites and tardigrades in Dominican amber

From time to time, small, fragile, previously unknown fossil invertebrates are found in specialized habitats. Occasionally, as in the present case, a fragment of the original habitat that existed millions of years ago is also preserved. The present article describes a previously unknown microinvertebrate in Dominican amber that cannot be placed in any group of extant invertebrates. Placed in a new family, genus, and species, the fossil shares characters with both tardigrades and mites, but clearly belongs to neither group. The several hundred fossil individuals preserved in the amber shared their moist, warm habitat with pseudoscorpions, nematodes, fungi, and protozoa. The large number of fossils provided additional evidence of their biology, including their reproductive behavior, developmental stages, and food. While there is no extant group that can accommodate these fossils, and we have no knowledge of any extant descendants, this discovery shows that unique lineages of minute invertebrates were surviving in the mid‐Tertiary.     

Carboniferous Onychophora from Montceau‐les‐Mines,France, and onychophoran terrestrialization

The geological age of the onychophoran crown‐group, and when the group came onto land, have been sources of debate. Although stem‐group Onychophora have been identified from as early as the Cambrian, the sparse record of terrestrial taxa from before the Cretaceous is subject to contradictory interpretations. A Late Carboniferous species from the Mazon Creek biota of the USA, Helenodora inopinata, originally interpreted as a crown‐group onychophoran, has recently been allied to early Cambrian stem‐group taxa. Here we describe a fossil species from the Late Carboniferous Montceau‐les‐Mines Lagerstätte, France, informally referred to as an onychophoran for more than 30 years. The onychophoran affinities of Antennipatus montceauensis gen. nov., sp. nov. are indicated by the form of the trunk plicae and the shape and spacing of their papillae, details of antennal annuli, and the presence of putative slime papillae. The poor preservation of several key systematic characters for extant Onychophora, however, prohibits the precise placement of the Carboniferous fossil in the stem or crown of the two extant families, or the onychophoran stem‐group as a whole. Nevertheless, A. montceauensis is the most compelling candidate to date for a terrestrial Paleozoic onychophoran.     

Soil and canopy Pseudoscorpiones (Arthropoda,Arachnida) in a monodominant forest of Attalea phalerata Mart. (Arecaceae) in the Brazilian Pantanal

We studied the occurrence of Pseudoscorpiones in the soil, leaf litter, and in canopies of a monodominant forest of Attalea phalerata at different seasons in the northern region of the Brazilian Pantanal. A total of 1197 pseudoscorpions from nine families and 16 species were sampled. Olpiidae, Chernetidae, and Geogarypidae predominated in soil and leaf litter. Chernetidae was the most abundant family in canopies. Soil and canopy corresponded to distinct habitats in relation to pseudoscorpion abundance and richness, with the canopies being the most diversified environment. These habitats are occupied in different ways by pseudoscorpion populations. Geogarypus sp. occurs in the edaphic environment during receding water and dry season, but can be found in canopies of A. phalerata exclusively during high water. This alternation in the use of the edaphic environments and canopies in the same area by pseudoscorpion species probably happens due to the strong seasonality of the Brazilian Pantanal.     

Rhabditophanes schneideri (Rhabditida) phoretic on a cave pseudoscorpion

Information of phoretic nematode-pseudoscorpion associations and cases of parasitism on five European species of pseudoscorpions was summarized by Cur?ic et al. [Curci?, B.P.M., Dimitrijevi?, R.N., Makarov, S.E., Luci?, L.R., Curci?, S.B., 1996. Further report on nematode-pseudoscorpion associations. Acta arachnol. 45, 43-46; Curci?, B.P.M., Sudhaus, W., Dimitrijevi?, R.N., Tomi?, V.T., Curci?, S.B., 2004. Phoresy of Rhabditophanes schneideri (Bütschli) (Rhabditida: Alloionematidae) on pseudoscorpions (Arachnida: Pseudoscorpiones). Nematology 6 (3), 313-317]. An examination of a sample of the cavernicolous pseudoscorpion Neobisium rajkodimitrijevici (Curci? and Tomi?, 2006) (comprising a holotype male and a paratype tritonymph) from a cave in Eastern Serbia revealed this false scorpion to be a nematode carrier; the present paper reports this finding and extends our knowledge of phoresy of Rhabditophanes on pseudoscorpions. This is the first time that the rhabditid R. schneideri (Bütschli, 1873) has been noted in association with a cavernicolous pseudoscorpion. There must be some patchily distributed micro-habitats in caves where saprobiotic nematodes and small arthropods can complete their life-cycles, for example something like deposits of bat guano. The transportation of Rhabditophanes J3 by pseudoscorpions indicate that Neobisium specimens often visit these micro-habitats to find their prey.     

A chiton without a foot

Abstract: The palaeoloricate ‘polyplacophorans’ are an extinct paraphyletic group of basal chiton‐like organisms known primarily from their fossilized valves. Their phylogenetic placement remains contentious, but they are likely to include both stem‐group Polyplacophora and stem‐group Aplacophora. Candidates for the latter position include ‘Helminthochiton’thraivensis from the Ordovician of Scotland, which we redescribe here through a combined optical and micro‐CT (XMT) restudy of the type material. The 11 specimens in the type series are all articulated, presenting partial or complete valve series as well as mouldic preservation of the girdle armature; they demonstrate a vermiform body plan. The valves are typically palaeoloricate in aspect, but differ in detail from all existing palaeoloricate genera; we hence erect Phthipodochiton gen. nov. to contain the species. The most notable feature of the fossils is the spicular girdle; this is impersistently preserved, but demonstrably wraps entirely around the ventral surface of the animal, implying that a ‘true’ (i.e. polyplacophoran like) foot was absent, although we do not exclude the possibility of a narrow solenogastre‐like median pedal groove having been present. Phthipodochiton thraivensis presents an apparent mosaic of aplacophoran and polyplacophoran features and as such will inform our understanding of the relationship between these groups of extant molluscs. An inference may also be drawn that at least some other palaeoloricates possessed an ‘armoured aplacophoran’ body plan, in contrast to the ‘limpet‐like’ body plan of extant Polyplacophora.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Olfactory organ of acipenseriformes and comparison with other actinopterygians: Patterns of diversity

The position and structure of the olfactory organ and its openings vary among actinopterygians. The anterior nasal opening is a simple perforation in the skin in many extant actinopterygians (e.g., acipenseriforms, lepisosteids, and primitive Recent teleosts) and represents the primitive condition. Polypterids and Amia each exhibit a derived condition, in which the anterior nasal opening extends into a tube. The olfactory organ is relatively far away from the anterior end of the elongate rostrum in acipenseriforms, whereas the olfactory organs are closer to the anterior end of the snout in extant actinopterygians (e.g., polypterids, lepisosteids, and amiids). In adults, olfactory organs are cuplike structures in most actinopterygians, but these organs are tubelike in polypterids. Among extant actinopterygians, a nasal diverticulum is present only in polypterids. Teleosts have accessory nasal sacs, but chondrosteans, polypterids, lepisosteids, and amiids lack them. The olfactory rosette is formed by primary folds or lamellae that may be placed anterior, lateral, posterior, and/or medial to the axis of the organ. Large acipenserids have 20–32 lamellae, polyodontids have 13–18 lamellae, lepisosteids have 8–10 lamellae, and Amia may have over 100. In teleosts, the number of lamellae varies from none or a few to over 200. Secondary lamellae are present in acipenseriforms, lepisosteids, and some advanced teleosts; secondary lamellae are interpreted as independently acquired in these lineages. Secondary lamellae are absent in Amia and primitive teleosts such as Elops and Hiodon. Tertiary lamellae are present in Acipenser oxyrhynchus. The arrangement of the primary lamellae in relation to the axis of the organ results in at least 11 patterns of the olfactory rosette in actinopterygians. Lamellae that are enclosed in a tubelike sac and that have an anteromedial diverticulum are specializations of polypterids. Primary lamellae anterior, lateral, and posterior to an elongate axis are characteristic of lepisosteids. The presence of primary lamellae lateral, medial, and posterior to an elongate olfactory axis is a synapomorphy of Halecomorpha (Amia plus teleosts). The absence of secondary lamellae is a synapomorphy of Halecomorpha. © 1994 Wiley-Liss, Inc.     

Making sense of ‘lower’ and ‘upper’ stem‐group Euarthropoda,with comments on the strict use of the name Arthropoda von Siebold, 1848

The ever‐increasing number of studies that address the origin and evolution of Euarthropoda – whose extant representatives include chelicerates, myriapods, crustaceans and hexapods – are gradually reaching a consensus with regard to the overall phylogenetic relationships of some of the earliest representatives of this phylum. The stem‐lineage of Euarthropoda includes numerous forms that reflect the major morphological transition from a lobopodian‐type to a completely arthrodized body organization. Several methods of classification that aim to reflect such a complex evolutionary history have been proposed as a consequence of this taxonomic diversity. Unfortunately, this has also led to a saturation of nomenclatural schemes, often in conflict with each other, some of which are incompatible with cladistic‐based methodologies. Here, I review the convoluted terminology associated with the classification of stem‐group Euarthropoda, and propose a synapomorphy‐based distinction that allows ‘lower stem‐Euarthropoda’ (e.g. lobopodians, radiodontans) to be separated from ‘upper stem‐Euarthropoda’ (e.g. fuxianhuiids, Cambrian bivalved forms) in terms of the structural organization of the head region and other aspects of overall body architecture. The step‐wise acquisition of morphological features associated with the origins of the crown‐group indicate that the node defining upper stem‐Euarthropoda is phylogenetically stable, and supported by numerous synapomorphic characters; these include the presence of a deutocerebral first appendage pair, multisegmented head region with one or more pairs of post‐ocular differentiated limbs, complete body arthrodization, posterior‐facing mouth associated with the hypostome/labrum complex, and post‐oral biramous arthropodized appendages. The name ‘Deuteropoda’ nov. is proposed for the scion (monophyletic group including the crown‐group and an extension of the stem‐group) that comprises upper stem‐Euarthropoda and Euarthropoda. A brief account of common terminological inaccuracies in recent palaeontological studies evinces the utility of Deuteropoda nov. as a reference point for discussing aspects of early euarthropod phylogeny.