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Concept: Chelicerata

9

Among the largest and most abundant aquatic predators during much of the early evolution of vertebrates, eurypterids have long been an iconic and intensely studied group of Paleozoic arthropods. We report a new specimen of the eurypterid Slimonia acuminata, which includes a fully articulated series of tail (postabdominal and telsonal) segments preserved in a tight lateral curve. Such a high degree of apparent lateral tail flexibility has not been previously recognized in eurypterids. From the perspective of hydrodynamics, the dorsoventrally flattened body plan of eurypterids would have limited the effectiveness of lateral tail motion as a means of propulsion. However, the long and serrated terminal tail spine of S. acuminata (and other eurypterids) would have made lateral tail strikes-which would have met a minimum of hydraulic resistance-an effective means of predatory attack and self-defense. Thus, many eurypterids are reinterpreted as substantially better-armed predators than previously supposed.

Concepts: Arthropod, Vertebrate, Scorpion, Chelicerata, Eurypterid, Eurypterida, Pterygotus, Slimonia

8

TO DATE, ONLY MORPHOLOGICAL AND ANATOMICAL DESCRIPTIONS OF MICROWHIP SCORPIONS (ARACHNIDA: Palpigradi) have been published. This very rare group is enigmatic not only in its relationships to other arachnids, but especially due to the fact that these animals dwell only underground (in caves, soil, and interstitial spaces). We observed the curious feeding habit of the microwhip scorpion Eukoenenia spelaea over the course of one year in Ardovsk√° Cave, located in Slovakia’s Karst region. We chose histology as our methodology in studying 17 specimens and based it upon Masson’s triple staining, fluorescent light and confocal microscopy. Single-celled cyanobacteria (blue-green algae) were conspicuously predominant in the gut of all studied palpigrades. Digestibility of the consumed cyanobacteria was supported by the presence of guanine crystals, glycogen deposits and haemocytes inside the palpigrade body. Cyanobacteria, the oldest cellular organisms on Earth, are very resistant to severe conditions in caves, including even darkness. Therefore, the cyanobacteria are able to survive in dark caves as nearly heterotrophic organisms and are consumed by cave palpigrades. Such feeding habit is extraordinary within the almost wholly predacious orders of the class Arachnida, and particularly so due to the type of food observed.

Concepts: Bacteria, Biology, Organism, Arachnid, Scorpion, Chelicerata, Arachnids, Palpigradi

7

Arachnids are a highly successful group of land-dwelling arthropods. They are major contributors to modern terrestrial ecosystems, and have a deep evolutionary history. Whip spiders (Arachnida, Amblypygi), are one of the smaller arachnid orders with ca. 190 living species. Here we restudy one of the oldest fossil representatives of the group, Graeophonus anglicus Pocock, 1911 from the Late Carboniferous (Duckmantian, ca. 315 Ma) British Middle Coal Measures of the West Midlands, UK. Using X-ray microtomography, our principal aim was to resolve details of the limbs and mouthparts which would allow us to test whether this fossil belongs in the extant, relict family Paracharontidae; represented today by a single, blind species Paracharon caecus Hansen, 1921.

Concepts: Evolution, Arthropod, Arachnid, Spider, Carboniferous, Chelicerata, Pedipalp, Amblypygi

6

As part of the German Barcode of Life campaign, over 3500 arachnid specimens have been collected and analyzed: ca. 3300 Araneae and 200 Opiliones, belonging to almost 600 species (median: 4 individuals/species). This covers about 60% of the spider fauna and more than 70% of the harvestmen fauna recorded for Germany. The overwhelming majority of species could be readily identified through DNA barcoding: median distances between closest species lay around 9% in spiders and 13% in harvestmen, while in 95% of the cases, intraspecific distances were below 2.5% and 8% respectively, with intraspecific medians at 0.3% and 0.2%. However, almost 20 spider species, most notably in the family Lycosidae, could not be separated through DNA barcoding (although many of them present discrete morphological differences). Conspicuously high interspecific distances were found in even more cases, hinting at cryptic species in some instances. A new program is presented: DiStats calculates the statistics needed to meet DNA barcode release criteria. Furthermore, new generic COI primers useful for a wide range of taxa (also other than arachnids) are introduced.

Concepts: Species, Arthropod, DNA barcoding, Arachnid, Spider, Chelicerata, Pedipalp, Arachnids

6

Spiders and scorpions are notorious for their fearful dispositions and their ability to inject venom into prey and predators, causing symptoms such as necrosis, paralysis, and excruciating pain. Information on venom composition and the toxins present in these species is growing due to an interest in using bioactive toxins from spiders and scorpions for drug discovery purposes and for solving crystal structures of membrane-embedded receptors. Additionally, the identification and isolation of a myriad of spider and scorpion toxins has allowed research within next generation antivenoms to progress at an increasingly faster pace. In this review, the current knowledge of spider and scorpion venoms is presented, followed by a discussion of all published biotechnological efforts within development of spider and scorpion antitoxins based on small molecules, antibodies and fragments thereof, and next generation immunization strategies. The increasing number of discovery and development efforts within this field may point towards an upcoming transition from serum-based antivenoms towards therapeutic solutions based on modern biotechnology.

Concepts: Arthropod, Arachnid, Toxin, Venom, Scorpion, Chelicerata, Spider bite, Pedipalp

6

MicroRNAs are small (~22 nucleotide) noncoding RNAs that repress translation and therefore regulate the production of proteins from specific target mRNAs. microRNAs have been found to function in diverse aspects of gene regulation within animal development and many other processes. Among invertebrates, both conserved and novel, lineage specific, microRNAs have been extensively studied predominantly in holometabolous insects such as Drosophila melanogaster However little is known about microRNA repertoires in other arthropod lineages such as the chelicerates. To understand the evolution of microRNAs in this poorly sampled subphylum, we characterized the microRNA repertoire expressed during embryogenesis of the common house spider Parasteatoda tepidariorum We identified a total of 148 microRNAs in P. tepidariorum representing 66 families. Approximately half of these microRNA families are conserved in other metazoans, while the remainder are specific to this spider. Of the 35 conserved microRNAs families 15 had at least two copies in the P. tepidariorum genome. A BLAST based approach revealed a similar pattern of duplication in other spiders and a scorpion, but not among other chelicerates and arthropods, with the exception of a horseshoe crab. Among the duplicated microRNAs we found examples of lineage specific tandem duplications, and the duplication of entire microRNA clusters in three spiders, a scorpion, and in a horseshoe crab. Furthermore, we found that paralogs of many P. tepidariorum microRNA families exhibit arm switching, which suggests that duplication was often followed by sub- or neofunctionalization. Our work shows that understanding the evolution of microRNAs in the chelicerates has great potential to provide insights into the process of microRNA duplication and divergence and the evolution of animal development.

Concepts: DNA, Gene, RNA, Insect, Arthropod, Arachnid, Spider, Chelicerata

6

Eurypterids are a group of extinct chelicerates that ranged for over 200 Myr from the Ordovician to the Permian. Gigantism is common in the group; about 50% of families include taxa over 0.8 m in length. Among these were the pterygotids (Pterygotidae), which reached lengths of over 2 m and were the largest arthropods that ever lived. They have been interpreted as highly mobile visual predators on the basis of their large size, enlarged, robust chelicerae and forward-facing compound eyes. Here, we test this interpretation by reconstructing the visual capability of Acutiramus cummingsi (Pterygotidae) and comparing it with that of the smaller Eurypterus sp. (Eurypteridae), which lacked enlarged chelicerae, and other arthropods of similar geologic age. In A. cummingsi, there is no area of lenses differentiated to provide increased visual acuity, and the interommatidial angles (IOA) do not fall within the range of high-level modern arthropod predators. Our results show that the visual acuity of A. cummingsi is poor compared with that of co-occurring Eurypterus sp. The ecological role of pterygotids may have been as predators on thin-shelled and soft-bodied prey, perhaps in low-light conditions or at night.

Concepts: Arthropod, Predation, Ecology, Eye, Spider, Trilobite, Chelicerata, Eurypterid

5

The public perception of spiders and scorpions is skewed towards the potential harm they can inflict in humans, despite recent scientific evidence that arachnid venom components might be useful as bioinsecticides or even human therapeutics. Nevertheless, arachnids are becoming more popular as pets in Europe, America and Asia, raising the question for regulatory agencies in these regions as to whether they need to take measurements to protect their citizens. In order to decide upon the necessary regulatory steps, they first need to determine which arachnids are actually dangerous to humans. This review therefore provides an overview of the current literature on verified bites and stings from spiders and scorpions with the aim of assessing their potential danger for human health. As a guideline, we also provide a list of those arachnid genera that we consider as potentially dangerous, which includes 10 spider and 11 scorpion genera. The arachnid genera classified as dangerous comprise less than a quarter of all extant scorpion species and only 0.5% of all spiders species, with the actual number most likely being much lower than that, as not all species in those genera might turn out to pose an actual threat for humans. In conclusion, we found that only a small percentage of scorpions and a minute percentage of all spiders can be considered as potentially dangerous to humans. While in some countries of origin the high incidence of envenomations by dangerous arachnids can result in a serious problem to the health system, we assessed the risk that the same species pose when kept as pets under controlled maintenance conditions as significantly lower.

Concepts: Species, Arthropod, Arachnid, Spider, Scorpion, Chelicerata, Pedipalp, Arachnids

5

The visual systems in chelicerates are poorly understood, even though they show strong variation in eye and visual neuropil architecture, thus may provide valuable insights for the understanding of chelicerate phylogeny and eye evolution. Comparable morphological characters are desperately sought for reconstructions of the phylogeny of Chelicerata, especially with respect to Arachnida. So far, reliable data exist only for Pycnogonida, Xiphosura, Scorpiones, and Araneae. The few earlier studies of the organisation of the visual system in harvestmen are contradictory concerning the number, morphology, and position of the visual neuropils.

Concepts: Arthropod, Arachnid, Spider, Scorpion, Chelicerata, Opiliones, Pedipalp, Chelicerae

5

A three-dimensionally preserved 2-mm-long larva of the arthropod Leanchoilia illecebrosa from the 520-million-year-old early Cambrian Chengjiang biota of China represents the first evidence, to our knowledge, of such an early developmental stage in a short-great-appendage (SGA) arthropod. The larva possesses a pair of three-fingered great appendages, a hypostome, and four pairs of well-developed biramous appendages. More posteriorly, a series of rudimentary limb Anlagen revealed by X-ray microcomputed tomography shows a gradient of decreasing differentiation toward the rear. This, and postembryonic segment addition at the putative growth zone, are features of late-stage metanauplii of eucrustaceans. L. illecebrosa and other SGA arthropods, however, are considered representative of early chelicerates or part of the stem lineage of all euarthropods. The larva of an early Cambrian SGA arthropod with a small number of anterior segments and their respective appendages suggests that posthatching segment addition occurred in the ancestor of Euarthropoda.

Concepts: Insect, Developmental biology, Arthropod, Crustacean, Annelid, Scorpion, Trilobite, Chelicerata