An unexpected diversity of Cyphophthalmi (Arachnida: Opiliones) in Upper Cretaceous Burmese amber.

Ten new Cyphophthalmi specimens (Arachnida: Opiliones) from the Upper Cretaceous (Lower Cenomanian) Burmese amber of northern Myanmar are described. Seven of these are placed in Stylocellidae, the predominant extant family found today in Southeast Asia. Sirocellus iunctus gen. et sp. nov. represents the first fossil with a combination of sironid and stylocellid characters, suggesting a still ongoing transition in some lineages during the Upper Cretaceous. Mesopsalis oblongus gen. et sp. nov. represents a second fossil with elongated ozophores, a character not known from modern species. Leptopsalis breyeri sp. nov. is the first Cretaceous cyphophthalmid assignable to an extant genus. The species Foveacorpus cretaceus gen. et sp. nov. and F. parvus gen. et sp. nov., which cannot be placed in an extant family, show morphological novelties for Cyphophthalmi such as numerous pits covering the whole body. The possible function of these pits is discussed. Three more adult males with unique adenostyles and two juveniles are not formally named but further indicate an already highly diverse cyphophthalmid fauna during the Cretaceous. The total number of named Burmese amber Cyphophthalmi species is raised from one to six, and the total fossil record for this suborder now stands at eight.

The "southeastern Europe" lineage of the brown dog tick Rhipicephalus sanguineus (sensu lato) identified as Rhipicephalus rutilus Koch, 1844: Comparison with holotype and generation of mitogenome reference from Israel.

The brown dog tick Rhipicephalus sanguineus (sensu lato) in the southeastern Mediterranean region and the Middle East is difficult to identify due to the presence of multiple mitochondrial DNA haplogroup lineages. The purpose of this study was to clarify the identity of the "southeastern Europe" lineage of this tick species complex. Our research shows that female ticks of the "southeastern Europe" lineage correspond to the morphology of R. rutilus Koch, 1844 as found in type-material at the Museum für Naturkunde Berlin in Germany. We characterised the complete mitogenomes of R. rutilus, R. turanicus Pomerantsev, 1940 and Rhipicephalus sanguineus (Latreille, 1806) in order to improve our understanding of the phylogenetic relationships among species within the R. sanguineus (sensu lato) complex. The material associated with the morphology of R. rutilus was previously labelled as the "southeastern Europe" lineage and found in Israel and Egypt, including Lower Egypt and the Nile Delta, where the original type-material was collected. Based on the morphology, genetic identity, and geographical distribution of the species, we conclude that the name R. rutilus is correctly linked to the "southeastern Europe" lineage of R. sanguineus (sensu lato).

Hard ticks in Burmese amber with Australasian affinities.

Three examples of metastriate hard ticks (Ixodida: Ixodidae) with apparent affinities to modern Australasian genera are described from the mid-Cretaceous (ca. 100 Ma) Burmese amber of Myanmar. Two nymphs of Bothriocroton muelleri sp. nov. represent the oldest (and only) fossil record of this genus, living members of which are restricted to Australia and predominantly feed on monitor lizards, snakes and echidnas. A female of Archaeocroton kaufmani sp. nov. shares its basis capitulum shape with the tuatara tick Archaeocroton sphenodonti (Dumbleton, 1943), the only extant member of this genus and an endemic species for New Zealand. The presence of 2 Australasian genera in Burmese amber is consistent with a previous record of an Ixodes Latreille, 1795 tick from this deposit which resembles Australian members of this genus. They further support an emerging hypothesis that fauna of the amber forest, which may have been on an island at the time of deposition, was at least partly Gondwanan in origin. A revised evolutionary tree for Ixodida is presented compiling data from several new Burmese amber ticks described in the last few years.

Harvestmen (Arachnida: Opiliones) in Eocene Rovno amber (Ukraine).

Six species of harvestman (Arachnida: Opiliones) are documented from the Eocene Rovno amber in Ukraine. From the suborder Eupnoi we record Caddo dentipalpus (C. L. Koch Berendt, 1854) (Caddidae), Amilenus deltshevi Dunlop Mitov, 2009 (Phalangiidae) and Dicranopalpus ramiger (C. L. Koch Berendt, 1854) (family incertae sedis). To these we add a new phalangiid, Metaphalangium martensi sp. n., which is the oldest representative of the Recent genus Metaphalangium Roewer, 1911 and new genus for the amber fauna. From Dyspnoi we record Sabacon claviger (Menge, 1854) (Sabaconidae) and propose Parahistricostoma gen. n. (Nemastomatidae), to accommodate Nemastoma tuberculatum C. L. Koch Berendt, 1854, yielding Parahistricostoma tuberculatum (C. L. Koch Berendt, 1854) comb. n. The Rovno harvestman fauna is briefly compared to both Baltic and Bitterfeld amber. The fossil species, C. dentipalpus, A. deltshevi, D. ramiger and P. tuberculatum occur in all three ambers, and S. claviger is found in Baltic and Rovno amber. The only genus and species unique to Rovno amber is thus our new taxon M. martensi. The composition of the Rovno opilionid fauna is discussed in the context of late Eocene palaeoenvironment and ecological preferences and the fossils are compared to the distribution of extant harvestman taxa. The oribatid mite Platyliodes ensigerus Sellnick, 1919 (Acari: Neoliodidae) is reported from Rovno amber for the first time.

The complete mitochondrial genome of the pentastomid Linguatula arctica (Pentastomida) from reindeer (Rangifer tarandus) in Northern Norway.

Here, we present the first complete mitochondrial genome of the pentastomid Linguatula arctica collected from the nasal passages of a reindeer (Rangifer tarandus) in Norway. The full length mitochondrial genome of L. arctica, which measures 14,789 bp in length, contains 13 protein-coding genes, 2 ribosomal RNA genes and 22 transfer RNA genes. A clear A + T bias is observed in the mitogenome of L. arctica with an overall base composition of 32.6% A, 27.5% T, 32.8% C, and 7,1% G., and a GC content of 39.9%. The gene arrangement is identical to that of previously described pentastomid mitogenomes.

Evolution: A Breath of Fresh Air for Eurypterids.

Eurypterids - Paleozoic 'sea scorpions' that could grow several meters long - were thought to have been aquatic. Computer tomography reveals that respiratory organs of a new fossil eurypterid resemble arachnid book lungs, supporting the hypothesis that eurypterids - and perhaps arachnid ancestors - were amphibious.

Trilobite compound eyes with crystalline cones and rhabdoms show mandibulate affinities.

Most knowledge about the structure, function, and evolution of early compound eyes is based on investigations in trilobites. However, these studies dealt mainly with the cuticular lenses and little was known about internal anatomy. Only recently some data on crystalline cones and retinula cells were reported for a Cambrian trilobite species. Here, we describe internal eye structures of two other trilobite genera. The Ordovician Asaphus sp. reveals preserved crystalline cones situated underneath the cuticular lenses. The same is true for the Devonian species Archegonus (Waribole) warsteinensis, which in addition shows the fine structure of the rhabdom in the retinula cells. These results suggest that an apposition eye with a crystalline cone is ancestral for Trilobita. The overall similarity of trilobite eyes to those of myriapods, crustaceans, and hexapods corroborates views of a phylogenetic position of trilobites in the stem lineage of Mandibulata.

Miniaturisation in Chelicerata.

Arachnids and their relatives (Chelicerata) range in body length from tens of centimetres in horseshoe crabs down to little more than 80-200 µm in several groups of mites. Spiders (Araneae) show the widest range within a given Bauplan - the largest species being ca. 270 times longer than the smallest - making them excellent models to investigate scaling effects. The two mite clades (Parasitiformes and Acariformes) are the main specialists in being small. Miniaturisation, and its consequences, is reviewed for both fossil and extant chelicerates. Morphological changes potentially related to miniaturisation, or adapting to the ecological niches that small size allows, include reduction in the length and number of legs, loss of prosomal arteries (and eventually also the heart), replacement of book lungs by tracheae, or even loss of all respiratory organs. There may also be evolutionary novelties, such as the acquisition of structures by which some mites attach themselves to larger hosts. The observed character distributions suggest a fairly fundamental division between larger pulmonate (lung-bearing) arachnids and smaller, non-pulmonate, groups which could reflect a phylogenetic dichotomy. However, it is worth noting that lineages of tiny spiders were originally fully pulmonate, but have acquired some typically non-pulmonate features, while camel spiders (Soli-fugae) can be large but have a Bauplan suggestive of smaller, non-pulmonate, ancestors.

Haemaphysalis cretacea a nymph of a new species of hard tick in Burmese amber.

The first fossil potentially assignable to the extant hard tick genus Haemaphysalis CL Koch (1844) (Ixodida: Ixodidae) is described from the Late Cretaceous (ca. 99 Ma) Burmese amber of Myanmar. Haemaphysalis (Alloceraea) cretacea sp. nov. is the oldest and only fossil representative of this genus; living members of which predominantly feed on mammals. Their typical hosts are known since at least the Jurassic and the discovery of a mid-Cretaceous parasite, which might have fed on mammals raises again the question of to what extent ticks are coupled to their (modern) host groups. An inferred Triassic split of Argasidae (soft ticks) into the bird-preferring Argasinae and mammal-preferring Ornithodorinae dates to about the time when dinosaurs (later including birds) and mammaliaforms as potential hosts were emerging. Ixodidae may have split into Prostriata and Metastriata shortly after the end-Permian mass extinction, an event which fundamentally altered the terrestrial vertebrate fauna. Prostriata (the genus Ixodes) prefer birds and mammals today, and some may have used groups like cynodonts in the Triassic. Basal metastriate ticks (e.g. Amblyomma) prefer reptiles, but derived metastriates (including Haemaphysalis) again prefer mammals. Here, we may be looking at a younger (Cretaceous?) shift associated with more recent mammalian radiations.

Microbial decay analysis challenges interpretation of putative organ systems in Cambrian fuxianhuiids.

The Chengjiang fossil Lagerstätte (Cambrian Stage 3) from Yunnan, southern China is renowned for its soft-tissue preservation. Accordingly structures in fuxianhuiids, radiodontans and great appendage arthropods have been interpreted as the nervous and cardiovascular systems, including brains, hearts and blood vessels. That such delicate organ systems survive the fossilization process seems remarkable; given that this mode of preservation involves major taphonomic changes, such as flattening, microbial degradation, chemical alteration and replacement. Here, we document a range of taphonomic preservation states in numerous articulated individuals of Fuxianhuia protensa We suggest that organic (partly iron mineral-replaced) bulbous structures in the head region, previously interpreted as brain tissue, along with sagittally located organic strands interpreted as part of the cardiovascular system or as nerve cords, may be better explained as microbial biofilms that developed following decomposition of the intestine, muscle and other connective tissues, forming halos surrounding the original organic remains.

Cretaceous arachnid Chimerarachne yingi gen. et sp. nov. illuminates spider origins.

Spiders (Araneae) are a hugely successful lineage with a long history. Details of their origins remain obscure, with little knowledge of their stem group and few insights into the sequence of character acquisition during spider evolution. Here, we describe Chimerarachne yingi gen. et sp. nov., a remarkable arachnid from the mid-Cretaceous (approximately 100 million years ago) Burmese amber of Myanmar, which documents a key transition stage in spider evolution. Like uraraneids, the two fossils available retain a segmented opisthosoma bearing a whip-like telson, but also preserve two traditional synapomorphies for Araneae: a male pedipalp modified for sperm transfer and well-defined spinnerets resembling those of modern mesothele spiders. This unique character combination resolves C. yingi within a clade including both Araneae and Uraraneida; however, its exact position relative to these orders is sensitive to different parameters of our phylogenetic analysis. Our new fossil most likely represents the earliest branch of the Araneae, and implies that there was a lineage of tailed spiders that presumably originated in the Palaeozoic and survived at least into the Cretaceous of Southeast Asia.

Terrestrial invertebrates in the Rhynie chert ecosystem.

The Early Devonian Rhynie and Windyfield cherts remain a key locality for understanding early life and ecology on land. They host the oldest unequivocal nematode worm (Nematoda), which may also offer the earliest evidence for herbivory via plant parasitism. The trigonotarbids (Arachnida: Trigonotarbida) preserve the oldest book lungs and were probably predators that practiced liquid feeding. The oldest mites (Arachnida: Acariformes) are represented by taxa which include mycophages and predators on nematodes today. The earliest harvestman (Arachnida: Opiliones) includes the first preserved tracheae, and male and female genitalia. Myriapods are represented by a scutigeromorph centipede (Chilopoda: Scutigeromorpha), probably a cursorial predator on the substrate, and a putative millipede (Diplopoda). The oldest springtails (Hexapoda: Collembola) were probably mycophages, and another hexapod of uncertain affinities preserves a gut infill of phytodebris. The first true insects (Hexapoda: Insecta) are represented by a species known from chewing (non-carnivorous?) mandibles. Coprolites also provide insights into diet, and we challenge previous assumptions that several taxa were spore-feeders. Rhynie appears to preserve a largely intact community of terrestrial animals, although some expected groups are absent. The known fossils are (ecologically) consistent with at least part of the fauna found around modern Icelandic hot springs.This article is part of a discussion meeting issue 'The Rhynie cherts: our earliest terrestrial ecosystem revisited'.

Amblyomma birmitum a new species of hard tick in Burmese amber.

Amblyomma birmitum sp. nov. is formally described as a new record from 99 Ma old Burmese amber from Myanmar. This confirms the presence of the extant hard tick genus Amblyomma C.L. Koch, 1844 (Ixodida: Ixodidae) in the Late Cretaceous. This discovery is placed in its wider context and some reports of fossil hard ticks, such as a Hyalomma C.L. Koch, 1844 in Eocene Baltic amber, are misidentifications. The genus Amblyomma belongs to the clade Metastriata, a group which probably also accommodates two extinct genera, Cornupalpatum Poinar and Brown, 2003 and Compluriscutata Poinar and Buckley, 2008, also found in Burmese amber. All three fossils are thus only a little younger than published molecular divergence time estimates (ca. 124 ± 17 Ma) for the Metastriata lineage. Amblyomma has a largely Gondwanan distribution today. However, in some biogeographical scenarios, e.g. the Samafrica model, its predicted radiation time postdates the dissolution of the original Gondwana supercontinent raising questions about how its current distribution pattern was achieved.

The phylogeny of fossil whip spiders.

BACKGROUND: 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. RESULTS: Tomography reveals several novel and significant character states for G. anglicus; most notably in the chelicerae, pedipalps and walking legs. These allowed it to be scored into a phylogenetic analysis together with the recently described Paracharonopsis cambayensis Engel & Grimaldi, 2014 from the Eocene (ca. 52 Ma) Cambay amber, and Kronocharon prendinii Engel & Grimaldi, 2014 from Cretaceous (ca. 99 Ma) Burmese amber. We recovered relationships of the form ((Graeophonus (Paracharonopsis + Paracharon)) + (Charinus (Stygophrynus (Kronocharon (Charon (Musicodamon + Paraphrynus)))))). This tree largely reflects Peter Weygoldt's 1996 classification with its basic split into Paleoamblypygi and Euamblypygi lineages; we were able to score several of his characters for the first time in fossils. Our analysis draws into question the monophyly of the family Charontidae. CONCLUSIONS: Our data suggest that Graeophonus is a crown group amblypygid, and falls within a monophyletic Paleoamblypgi clade, but outside the family Paracharontidae (= Paracharonopsis + Paracharon). Our results also suggest a new placement for the Burmese amber genus Kronocharon, a node further down from its original position. Overall, we offer a broad phylogenetic framework for both the fossil and Recent whip spiders against which future discoveries can be tested.

Segmentation and tagmosis in Chelicerata.

Patterns of segmentation and tagmosis are reviewed for Chelicerata. Depending on the outgroup, chelicerate origins are either among taxa with an anterior tagma of six somites, or taxa in which the appendages of somite I became increasingly raptorial. All Chelicerata have appendage I as a chelate or clasp-knife chelicera. The basic trend has obviously been to consolidate food-gathering and walking limbs as a prosoma and respiratory appendages on the opisthosoma. However, the boundary of the prosoma is debatable in that some taxa have functionally incorporated somite VII and/or its appendages into the prosoma. Euchelicerata can be defined on having plate-like opisthosomal appendages, further modified within Arachnida. Total somite counts for Chelicerata range from a maximum of nineteen in groups like Scorpiones and the extinct Eurypterida down to seven in modern Pycnogonida. Mites may also show reduced somite counts, but reconstructing segmentation in these animals remains challenging. Several innovations relating to tagmosis or the appendages borne on particular somites are summarised here as putative apomorphies of individual higher taxa. We also present our observations within the concept of pseudotagma, whereby the true tagmata - the prosoma and opisthosoma - can be defined on a fundamental change in the limb series while pseudotagmata, such as the cephalosoma/proterosoma, are expressed as divisions in sclerites covering the body without an accompanying change in the appendages.

The complete mitochondrial genome of the pentastomid Armillifer grandis (Pentastomida) from the Democratic Republic of Congo.

We present the first complete mitochondrial genome of the pentastomid Armillifer grandis (Arthropoda: Pentastomida) collected from the lungs of a rhinoceros viper (Bitis nasicornis) in the Democratic Republic of Congo. The full length mitochondrial genome of Armillifer grandis, which measures 16,073 bp in length, contains 13 protein-coding genes, 2 ribosomal RNA genes, and 22 transfer RNA genes. A clear A + T bias is observed in the mitogenome of Armillifer grandis with an overall base composition of 34.6% A, 29.4% T, 29% C, and 6.9% G, and a GC content of 35.9%. The gene arrangement is identical to that of previously described pentastomid mitogenomes.

Microtomography of the Baltic amber tick Ixodes succineus reveals affinities with the modern Asian disease vector Ixodes ovatus.

BACKGROUND: Fossil ticks are extremely rare and Ixodes succineus Weidner, 1964 from Eocene (ca. 44-49 Ma) Baltic amber is one of the oldest examples of a living hard tick genus (Ixodida: Ixodidae). Previous work suggested it was most closely related to the modern and widespread European sheep tick Ixodes ricinus (Linneaus, 1758). RESULTS: Restudy using phase contrast synchrotron x-ray tomography yielded images of exceptional quality. These confirm the fossil's referral to Ixodes Latreille, 1795, but the characters resolved here suggest instead affinities with the Asian subgenus Partipalpiger Hoogstraal et al., 1973 and its single living (and medically significant) species Ixodes ovatus Neumann, 1899. We redescribe the amber fossil here as Ixodes (Partipalpiger) succineus. CONCLUSIONS: Our data suggest that Ixodes ricinus is unlikely to be directly derived from Weidner's amber species, but instead reveals that the Partipalpiger lineage was originally more widely distributed across the northern hemisphere. The closeness of Ixodes (P.) succineus to a living vector of a wide range of pathogens offers the potential to correlate its spatial and temporal position (northern Europe, nearly 50 million years ago) with the estimated origination dates of various tick-borne diseases.