168 million years old "marine lice" and the evolution of parasitism within isopods.

BACKGROUND: Isopods (woodlice, slaters and their relatives) are common crustaceans and abundant in numerous habitats. They employ a variety of lifestyles including free-living scavengers and predators but also obligate parasites. This modern-day variability of lifestyles is not reflected in isopod fossils so far, mostly as the life habits of many fossil isopods are still unclear. A rather common group of fossil isopods is Urda (190-100 million years). Although some of the specimens of different species of Urda are considered well preserved, crucial characters for the interpretation of their lifestyle (and also of their phylogenetic position), have so far not been accessible. RESULTS: Using up-to-date imaging methods, we here present morphological details of the mouthparts and the thoracopods of 168 million years old specimens of Urda rostrata. Mouthparts are of a sucking-piercing-type morphology, similar to the mouthparts of representatives of ectoparasitic isopods in groups such as Aegidae or Cymothoidae. The thoracopods bear strong, curved dactyli most likely for attaching to a host. Therefore, mouthpart and thoracopod morphology indicate a parasitic lifestyle of Urda rostrata. Based on morphological details, Urda seems deeply nested within the parasitic isopods of the group Cymothoida. CONCLUSIONS: Similarities to Aegidae and Cymothoidae are interpreted as ancestral characters; Urda is more closely related to Gnathiidae, which is therefore also interpreted as an ingroup of Cymothoida. With this position Urda provides crucial information for our understanding of the evolution of parasitism within isopods. Finally, the specimens reported herein represent the oldest parasitic isopods known to date.

Diversity and distribution patterns of the Oligocene and Miocene decapod crustaceans (Crustacea: Malacostraca) of the Western and Central Paratethys.

Decapod associations have been significant components of marine habitats throughout the Cenozoic when the major diversification of the group occurred. In this respect, the circum-Mediterranean area is of particular interest due to its complex palaeogeographic history. During the Oligo-Miocene, it was divided in two major areas, Mediterranean and Paratethys. Decapod crustaceans from the Paratethys Sea have been reported in the literature since the 19th century, but only recent research advances allow evaluation of the diversity and distribution patterns of the group. Altogether 176 species-level taxa have been identified from the Oligocene and Miocene of the Western and Central Paratethys. Using the three-dimensional NMDS analysis, the composition of decapod crustacean faunas of the Paratethys shows significant differences through time. The Ottnangian and Karpatian decapod associations were similar to each other both taxonomically and in the mode of preservation, and they differed taxonomically from the Badenian ones. The Early Badenian assemblages also differed taxonomically from the Late Badenian ones. The time factor, including speciation, immigration from other provinces and/or (local or global) extinction, can explain temporal differences among assemblages within the same environment. High decapod diversity during the Badenian was correlated with the presence of reefal settings. The Badenian was the time with the highest decapod diversity, which can, however, be a consequence of undersampling of other time slices. Whereas the Ottnangian and Karpatian decapod assemblages are preserved virtually exclusively in the siliciclastic "Schlier"-type facies that originated in non-reefal offshore environments, carbonate sedimentation and the presence of reefal environments during the Badenian in the Central Paratethys promoted thriving of more diverse reef-associated assemblages. In general, Paratethyan decapods exhibited homogeneous distribution during the Oligo-Miocene among the basins in the Paratethys. Based on the co-occurrence of certain decapod species, migration between the Paratethys and the North Sea during the Early Miocene probably occurred via the Rhine Graben. At larger spatial scales, our results suggest that the circum-Mediterranean marine decapod taxa migrated in an easterly direction during the Oligocene and/or Miocene, establishing present-day decapod communities in the Indo-West Pacific.

Euryhaline preferences of the decapod crab Mioplax socialis enabled it to survive during the Badenian/Sarmatian extinction (Miocene) in the Central Paratethys.

Although decapod crustaceans of the Central Paratethys were diverse during the Badenian (Langhian-Early Serravallian), a dramatic drop in their diversity occurred at the boundary with the Sarmatian. A crab Mioplax socialis is one of the few decapods reported from the Lower Sarmatian (Mohrensternia Zone) of the Paratethys. Until now, this species has been known from only a handful of specimens from Austria, Croatia and Bulgaria (Central Paratethys), and its systematics and ecology remain poorly known. Here, on the basis of new specimens from the Sarmatian tuffitic clays of the Stretava Formation (Skáros, Eastern Slovakia) we confirm that this species belongs to the subfamily Chasmocarcininae. The diagnostic characters of the male sternum that allow this classification are reported for the first time. The molluscan assemblage co-occurring with M. socialis demonstrate that this species tolerated conditions with variable salinity. Its tolerance of a broad range of salinity regimes may thus explain its survival across the Badenian-Sarmatian extinction event. Preservation of near-complete and fully articulated individuals of M. socialis suggests calm conditions and short residence times on the sediment-water interface.

Ghost shrimps (Decapoda: Axiidea: Callianassidae) as producers of an Upper Miocene trace fossil association from sublittoral deposits of Lake Pannon (Vienna Basin, Slovakia).

Numerous trace fossils are described from the Late Miocene sediments of the Bzenec Formation exposed at the Gbely section (the Vienna Basin, Slovakia). During deposition of the sediments the area was part of the large, long-lived brackish to freshwater Lake Pannon. Most of the trace fossils are attributed herein to Egbellichnus jordidegiberti igen et ispec. nov. and are interpreted as burrows produced by decapod crustaceans, specifically by a ghost shrimp of the family Callianassidae. This interpretation is based on two independent lines of evidence: environmental requirements of large bioturbators and the burrow morphology itself. The new ichnotaxon is distinguished from other related ichnotaxa by a combination of typically inclined (roughly at an angle of 45°) cylindrical burrows, absence of lining, and tunnels making loops or bends at approximately right angles. The burrow systems at Gbely document the survival of ghost shrimp long after the closure of all seaways and the origin of Lake Pannon. As today, no ghost shrimp are known from long-lived brackish lakes. Egbellichnus from Gbely is the only, although indirect, record of ghost shrimp from a brackish lake environment reported so far.

Decapod Crustacea of the Central Paratethyan Ottnangian Stage (middle Burdigalian): implications for systematics and biogeography.

Decapod crustaceans from the Ottnangian (middle Burdigalian, Lower Miocene) of the Western and Central Paratethys remain poorly known. In this study, we review and re-describe mud shrimps (Jaxea kuemeli), ghost shrimps (Gourretia sp., Calliax michelottii) and brachyuran crabs of the families Leucosiidae, Polybiidae and Portunidae. A dorsal carapace of the genus Calliax is reported for the first time in the fossil record. Re-examination of the type material of Randallia strouhali (Leucosiidae) and Geryon ottnangensis (Geryonidae) resulted in a transfer of these species into Palaeomyra (Leucosiidae) and Liocarcinus (Polybiidae), respectively. Achelous vindobonensis, originally described as a chela of a portunid crab, probably belongs to a member of Polybiidae and is provisionally treated as Liocarcinus sp. Only two species, J. kuemeli and C. michelottii, are also known from the Karpatian, the succeeding Paratethyan stage. In most cases, the decapod assemblages of the Ottnangian consist of rather shallow-water taxa whereas the assemblages of the Karpatian consist of deep-water taxa from the middle and outer shelf. The Central Paratethyan assemblages show similarities in genus composition to the Proto-Mediterranean and recent Indo-Pacific regions. Gourretia sp. represents the earliest occurrence of the respective genus in the fossil record. The Oligocene-Early Miocene appearance of Palaeomyra and Liocarcinus in the circum-Mediterranean implies that sources of present-day diversity hotspots in the Indo-Pacific trace to the Western Tethys (as for other decapod genera), although coeval decapod assemblages in the Indo-Pacific remain poorly known.

Deep-water fossorial shrimps from the Oligocene Kiscell Clay of Hungary: Taxonomy and palaeoecology.

We describe deep-water ghost shrimp assemblages from the otherwise well known Oligocene Kiscell Clay in Hungary. The described fossorial shrimps (Decapoda: Callianassidae and Ctenochelidae) include: Ctenocheles rupeliensis (younger synonym Callianassa nuda) and Lepidophthalmus crateriferus (younger synonym Callianassa brevimanus). The fossil material of the former species is assigned to Ctenocheles based on the morphology of the major cheliped, particularly the pectinate fingers, bulbous propodus, cup-shaped carpus and elongated merus. Lepidophthalmus crateriferus from the Oligocene of Hungary is the first unequivocal fossil record of the genus, which is distinguished in the fossil record on the basis of the presence of a meral blade and meral hook on the major cheliped. Lepidophthalmus is today known exclusively from shallow-water environments. The finding of a deep-water fossil representative of Lepidophthalmus therefore appears to be a reverse of the common pattern of groups shifting environments from onshore to offshore over geological time, as seen in many taxa. The presence of Lepidophthalmus crateriferus comb. nov. in the Kiscell Clay therefore suggests different ecological requirements for at least some populations of this genus in the geological past.

On the occurrence of Ctenocheles (Decapoda, Axiidea, Ctenochelidae) in the Bohemian Cretaceous Basin.

Because of close morphological affinities, fossil cheliped fragments of the ghost shrimp Ctenocheles (Decapoda, Axiidea, Ctenochelidae) can be easily misidentified as remains of different decapod crustacean taxa. Re-examination of the Cretaceous decapods deposited in the National Museum in Prague revealed that all supposed specimens of the lobster genus Oncopareia found in the Middle Coniacian calcareous claystones of the Brezno Formation, including one of the Fritsch's original specimens of Stenocheles parvulus, actually belong to Ctenocheles. This material together with newly collected specimens from the same locality, allowed for erection of a new species, Ctenocheles fritschi. Its major chela possesses a serrated ischium and ovoid, unarmed merus; therefore, it is considered a close relative of the extant C. collini and C. maorianus. Ctenocheles fritschi sp. nov. represents the first report on the occurrence of the genus from the Bohemian Cretaceous Basin. It is one of the oldest records of Ctenocheles and simultaneously one of the best preserved fossils of the genus reported to date. Confusing taxonomy of S. parvulus is reviewed and shortly discussed.

Revision of the Miocene shrimp Callianassa kerepesiensis Müller, 1976 (Malacostraca, Decapoda), with a description of a new species.

Based on the re-examination of the type and additional material of a fossorial shrimp Callianassa kerepesiensis Müller, 1976, from the middle Miocene (Badenian) of Hungary, the species is re-assigned to Calliax de Saint Laurent, 1973. Additionally, Callianassa reinhardpalorum sp. nov. is described based on the morphologically similar cheliped elements, classified at one time as C. cf. kerepesiensis. Both species, Callianassa reinhardpalorum sp. nov. and Calliax kerepesiensis comb. nov., have extant congeners inhabiting modern Mediterranean Sea and north-east Atlantic. The sedimentology of the localities with remains of the studied fossil taxa corresponds with the environmental preferences of their extant congeners, suggesting they inhabited marine settings with slightly changing salinity and the water depth not exceeding 20 m.

A new species of Bathynomus A. Milne-Edwards, 1879 (Malacostraca: Peracarida: Isopoda: Cirolanidae) from the upper Miocene of the Guadalquivir Basin (Spain).

Today representatives of the giant cirolanid isopods of the genus Bathynomus inhabit tropical to warm-temperate waters of the West Atlantic and the Indo-West Pacific. In not so distant geological past, however, the genus was present also in the Mediterranean, as documented by numerous fossils from the Plio-Pleistocene of Italy. Herein, a somewhat older occurrence of the genus is reported, from the North-East Atlantic. Based on a single posterior moult from the upper Miocene (Tortonian) of southwestern Spain, a new species is described. Bathynomus civisi sp. nov. differs from its congeners by both uropod rami having the obtuse distolateral corner produced as a distinct tip and a wider gap between the second and third side distal spines than between others. The new species is so far known only from the type locality in close vicinity of Arroyo Trujillo, in the municipality of Cantillana (Seville, Spain), where the marls of the Écija Formation are exposed. At the time of deposition of these marls, the area represented a part of the Norbetic Strait, a corridor connecting the North-East Atlantic with the Mediterranean. The new occurrence reported herein further supports the environmental preferences of the genus throughout its geological past for deep-marine settings.

The oldest freshwater crabs: claws on dinosaur bones.

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochthony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of "true" crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.

Comprehensive analysis and reinterpretation of Cenozoic mesofossils reveals ancient origin of the snapping claw of alpheid shrimps.

Alpheid snapping shrimps (Decapoda: Caridea: Alpheidae) constitute one of the model groups for inferences aimed at understanding the evolution of complex structural, behavioural, and ecological traits among benthic marine invertebrates. Despite being a super-diverse taxon with a broad geographical distribution, the alpheid fossil record is still poorly known. However, data presented herein show that the strongly calcified fingertips of alpheid snapping claws are not uncommon in the fossil record and should be considered a novel type of mesofossil. The Cenozoic remains analysed here represent a compelling structural match with extant species of Alpheus. Based on the presence of several distinct snapping claw-fingertip morphotypes, the major radiation of Alpheus lineages is estimated to have occurred as early as 18 mya. In addition, the oldest fossil record of alpheids in general can now be confirmed for the Late Oligocene (27-28 mya), thus providing a novel minimum age for the entire group as well as the first reliable calibration point for deep phylogenetic inferences.

Balsscallichirus Sakai, 2011 (Decapoda: Axiidea: Callianassidae) in the fossil record: systematics and palaeobiogeography.

The fossil record of the ghost shrimp genus Balsscallichirus Sakai, 2011 (Crustacea: Decapoda: Callianassidae) is revised. Barnardcallichirus Sakai, 2011 and Tirmizicallichirus Sakai, 2011 are considered subjective synonyms of Balsscallichirus. Based on the examination of extant species it is argued that the morphology of the major cheliped merus, in combination with other hard part morphology characters, is sufficient for assignment of the fossil material into the genus. Main identifying characters are on merus which is keeled along its midline and its lower half is tuberculated; its lower margin possesses broad proximal meral hook continuing into a lobe distally; the entire lower margin is subdivided into numerous irregularly spaced spines. Three species, Callianassa sismondai A. Milne-Edwards, 1860, C. floriana Glaessner, 1928, and Podocallichirus laepaensis Hyzný & Muñiz, 2012, originally described from the Miocene of Italy, Austria and Spain, respectively, are assigned to Balsscallichirus herein. Neocallichirus wellsi Schweitzer, Feldmann & Gingerich, 2004 from the Upper Eocene of Pakistan is tentatively assigned to that genus as well. Spatial and temporal distribution of the genus indicates that at least since the Oligocene, and possibly even sooner (the Late Eocene), the genus has been restricted to the Western Tethys Region. Later, it migrated also into West Atlantic establishing present day communities.

A new iphiculid crab (Crustacea, Brachyura, Leucosioidea) from the Middle Miocene of Austria, with notes on palaeobiogeography of Iphiculus.

A new fossil species of the iphiculid genus Iphiculus Adams & White, 1849, (Crustacea, Brachyura) is described on the basis of three specimens from the Middle Miocene Florian Beds of Styria, Austria. Iphiculus eliasi sp. nov. constitutes the first European record of the genus. This occurrence represents the oldest record of Iphiculus, having implications for the palaeobiogeographic history of the family Iphiculidae. It is suggested that Iphiculus may have originated in the Western Tethys and migrated subsequently into the Indo-West Pacific. Alternatively, its current geographic restriction to the Indo-West Pacific can be a remnant of an ancient broader geographic distribution.

From the Palaeontological Collection of the Universalmuseum Joanneum - The Cenozoic Decapod Crustaceans (Crustacea: Malacostraca: Decapoda).

Cenozoic decapod crustaceans housed in the collections of the Universalmuseum Joanneum (Graz, Austria) are reviewed. Previous descriptions, geographic and stratigraphic provenance and collection history are discussed. Altogether 72 specimens are figured, including five holotypes. Taxonomic affinity of previously unpublished material is addressed. Gebiacantha sp. from the middle Miocene of Wetzelsdorf is the first fossil record of the genus from the Paratethys.

A catalogue of the type and figured fossil decapod crustaceans in the collections of the Geological Survey of Austria in Vienna.

The present catalogue lists and figures the type and figured material of fossil decapod crustaceans housed in the collections of the Geological Survey of Austria in Vienna. Specimens previously believed to be lost were relocated. Lectotypes and paralectotypes are chosen herein for 11 species. Taxonomic affinity of taxa originally described in open nomenclature is discussed.

An appraisal of Typilobus Stoliczka, 1871 (Crustacea, Brachyura, Leucosioidea), with description of a new family and genus.

Since its description, Typilobus Stoliczka, 1871, has become something like a catch-all taxon to accommodate fossil leucosioid crabs with uncertain affinities. As many as 17 species have been assigned to Typilobus, recorded from the Eocene to Miocene strata of Europe, Africa, and Asia. The re-examination of the type species, Typilobus granulosus Stoliczka, 1871, from the Miocene of Pakistan, has shown inconsistency in its description and published figures leading to the need of a revision of all species assigned to Typilobus. Typilobus boscoi Vía Boada, 1959, from the Middle Eocene of Spain, is revised and assigned to Folguerolesia n. gen. Characters present on ventral portions of male and female specimens differ from all known representatives of Leucosioidea, although the general morphology supports inclusion in this superfamily. A new family, Folguerolesiidae n. fam. is thus erected to accommodate Folguerolesia boscoi comb. nov., leading to recognition of three families of leucosioid crabs, including Iphiculidae and Leucosiidae.

New Early Cenozoic ghost shrimps (Decapoda, Axiidea, Callianassidae) from Pakistan and their palaeobiogeographic implications.

A new set of Paleocene and Eocene decapod crustaceans is described from the Kirthar Range of Pakistan. Two new ghost shrimps (Crustacea, Decapoda, Callianassidae) are described: Neocallichirus khadroensis Hyzný & Charbonnier, n. sp. from the Paleocene (Danian, Khadro Formation) of Gawar Band, Ranikot District, and Neocallichirus lakhraensis Hyzný & Charbonnier, n. sp. from the Early Eocene (Ypresian, Lakhra Formation) of Rbod Nala, Jhirak District. Both new species exhibit chelipeds which are morphologically surprisingly close to extant Neocallichirus karumba (Poore & Griffin, 1979) from the Indo-West Pacific. A group of species sharing this same cheliped morphology is provisionally called the "karumba group" based on Neocallichirus karumba, best documented species. The "karumba group" encompasses seven fossil species: the two new Pakistani species, Neocallichirus tuberculatus (Lorenthey in Lorenthey & Beurlen, 1929) n. comb. from the Eocene of Hungary, Neocallichirus borensis Beschin, De Angeli, Checchi & Mietto, 2006 from the Eocene of Italy, Neocallichirus birmanicus (Noetling, 1901) n. comb. from the Miocene of Myanmar, Neocallichirus dijki (Martin, 1883) from the Miocene of Java and Philippines, and the subfossil Neocallichirus maximus (A. Milne-Edwards, 1870) from Thailand. Based upon the extant and fossil occurrences, it is difficult to reconstruct migration pattern of the "karumba group". For now, it can be concluded, that at the genus level, a relative homogeneity of the ghost shrimps is observed between the Eastern and the Western Tethyan regions, as already suggested by Merle et al. (2014) for the assemblage of volutid gastropods from the Lakhra Formation.

Barremian decapod crustaceans from Serre de Bleyton (Drôme, SE France).

Based on mostly small-sized isolated cheliped fingers, a new decapod crustacean assemblage is described from the Barremian of Serre de Bleyton (Drôme, SE France). The assemblage is composed mostly of representatives of the crab family Dynomenidae. In addition, remains of astacidean lobsters, axiidean shrimps, paguroid hermit crabs and brachyurous crabs of the families Necrocarcinidae and ?Cenomanocarcinidae occur in low numbers. Graptocarcinus moosleitneri (Dynomenidae) and ?Paranecrocarcinus schloegli (Necrocarcinidae) are introduced as new species. They both exhibit presence of multi-setal pores on dactyli that are interpreted as parts of a sieving mechanism used in feeding. The stratigraphic range of Graptocarcinus is extended herein to the Barremian.