First Fossil Xiphosuran (Chelicerata,Xiphosura) Egg Clutch from the Carboniferous of Khakassia

A new formal genus with a new species Xiphosuroides khakassicus gen. et sp. nov. is established for a clutch of nine eggs containing embryos, presumably xiphosurans (Chelicerata, Xiphosura, Bellinurina). This unique specimen comes from the Lower–Middle Carboniferous Sarskaya Formation of Khakassia (Izykhskie Kopi locality). If the hypothesis of the xiphosuran nature of these eggs is correct, the specimen described in this paper is the first certain record of fossil xiphosuran eggs and embryos.     

Xiphosuran Digging Traces at the Late Carboniferous Joggins Fossil Cliffs UNESCO World Heritage Site,Nova Scotia,Canada

One hundred Selenichnites rossendalensis (excavations), sometimes with associated trackways, have been found at Joggins Fossil Cliffs (Nova Scotia, Canada; Pennsylvanian). The majority of the Selenichnites found in this study contain only one lunate cast and some are distinguished by linear ridges that project behind a single lunate cast. A detailed investigation of the morphology and orientation of these Selenichnites—attributable to xiphosurans, relatives of modern horseshoe crabs (Limulus polyphemus)—yields an understanding of xiphosuran digging behavior. Statistical analysis of 96 lunate casts from two populations of Selenichnites separated by several hundred thousand years shows, with 99% confidence, that the prosomas (heads) of the younger population are smaller. Further study of the wealth of Selenichnites at Joggins will be useful to determine which factors control xiphosuran body-size changes and the timescale on which those changes can be seen manifest in a population. The smallest known Selenichnites (7 mm wide) is also reported.     

A CARBONIFEROUS SYNZIPHOSURINE (XIPHOSURA) FROM THE BEAR GULCH LIMESTONE, MONTANA, USA

Abstract:  A new synziphosurine, Anderella parva gen. et sp. nov., extends the known range of this group from the Silurian to the Carboniferous and is the youngest known so far from the fossil record. Previously the youngest synziphosurine, Kasibelinurus , was from the Devonian of North America. Anderella parva has a semi-oval carapace with pointed genal regions, nine freely articulating opisthosomal segments and a long styliform tail spine. It is the third xiphosuran genus to be described from the Bear Gulch Limestone and its discovery highlights this deposit as containing an unusually diverse xiphosuran biota. It is also the only known locality where synziphosurines occur alongside the more derived xiphosurids. Xiphosurans reached their greatest diversity in the Carboniferous when the xiphosurids began to occupy brackish and freshwater habitats and became dominant over the synziphosurines. The discovery of the only known Carboniferous synziphosurine in marine sediments may indicate their inability to exploit these same environments.     

Changed number of the collection of Permian ammonoids from Novaya Zemlya housed in the cited collection repository of the Borissiak Paleontological Institute,Russian Academy of Sciences

The development of views on the phylogeny of the family Cyclolobidae Zittel, 1895 is discussed. The generic composition is emended, phylogenetic links are indicated, a new version of the phylogenetic scheme is proposed, and phylogeny of the key genera is traced at species level. The sutural ontogeny of the genus Mexicoceras is examined, its placement in the family Cyclolobidae is supported, and its position in the phylogenetic reconstruction of the family is determined.     

The Ichnogenus Kouphichnium and Related Xiphosuran Traces from the Steven C. Minkin Paleozoic Footprint Site (Union Chapel Mine), Alabama,USA: Ichnotaxonomic and Paleoenvironmental Implications

Abstract

The ichnogenus Kouphichnium and associated ichnofossils attributed to xiphosuran activity are here re-examined from samples collected from the Pennsylvanian-age Steven C. Minkin Fossil Site at the Union Chapel Mine, in Walker County, Alabama, USA. The large sample size offers an unique opportunity to evaluate some Kouphichnium ichnospecies. Thus, the morphological variability resulting from the taphonomic, ethological, taxonomic variability and underprint fallout are evaluated using this large sample set. Three morphotypes have been segregated from the material previously assigned to K. aspodon discovered at this and adjacent sites. Ichnospecies of Kouphichnium identified at the Union Chapel Mine site include: K. lithographicum, K. aspodon and two new ichnospecies (K. atkinsoni and K. minkinensis). Additionally, Kouphichnium-like traces that are associated with “jumper” traces have been previously misinterpreted as Kouphichnium and Selenichnites, respectively, and are excluded from this study, leaving them in open nomenclature as they will be the subject of a subsequent publication. We here redescribe the holotype of K. aspodon and designate lectoparatypes to better define the ichnospecies. New trace makers for some Kouphichnium ichnospecies are hypothesized, in contrast to the traditional xiphosuran attribution.     

Revision of the Permian ammonoid family Cyclolobidae

The development of views on the phylogeny of the family Cyclolobidae Zittel, 1895 is discussed. The generic composition is emended, phylogenetic links are indicated, a new version of the phylogenetic scheme is proposed, and phylogeny of the key genera is traced at species level. The sutural ontogeny of the genus Mexicoceras is examined, its placement in the family Cyclolobidae is supported, and its position in the phylogenetic reconstruction of the family is determined.     

Révision taxonomique du genre d'ammonite téthysien Simoceras et création d'un nouveau genre Volanoceras (Ammonitina,Simoceratidae)

The type-species of the genus SimocerasZittel isS. biruncinatum (Qu.). This genus includes the species-group biruncinatum (Qu.) — admirandum (Zitt.). The new genus Volanoceras is created for the volanense (Opp.) species-group which is excluded from the genus Simoceras.     

The Nymphal Eyes of Parabuthus transvaalicus Purcell, 1899 (Buthidae): An Accessory Lateral Eye in a Scorpion

In P. transvaalicus nymphs, 5 pairs of lateral ocelli each composed of a corneal lens, R-cell units forming a latticed rhabdom, arhabdomeric cells and pigment cells are present. In addition, we found a pair of unpigmented accessory sense organs situated ventroposteriorly to the lateral ocelli in prenymphs as well as in first nymphs. They are composed of primary, rhabdomeric sensory cells, and we infer that they represent a second type of lateral eye. They also comprise sensory units, but lenses and screening pigment are lacking. Their position and cellular architecture corresponds well with that of the “rudimentary” lateral eye of the xiphosuran, Limulus. The occurrence of a bipartite lateral visual system in Chelicerata and Arthropoda is discussed.     

Unusual skeletal morphology and systematic description of a new Devonian cryptostome bryozoan from the Western Sahara

Palaeozoic bryozoan colonies display a large variety of skeletal elements. Various types of rod-like styles are mainly built by laminated and non-laminated (hyaline) skeleton. Typical styles consist of hyaline cores and surrounding laminated sheaths. They usually protrude on the colony surface as spines. In a new bryozoan genus from the Middle Devonian of the Western Sahara described here, styles do not protrude on the colony surface and are embedded within the laminated skeleton. They consist of fibrous material. This new type of styles, unknown from other bryozoans, is here called a “cryptostyle”. A special characteristic of these structures is their intensive reddish-brown colouration. This colouration is apparently caused by the presence of ferric iron between the individual fibres. The function of cryptostyles was apparently weight reduction and stabilising the skeleton. The general morphology of the new bryozoan, Cryptostyloecia hexapuncta gen. et n. sp., implies its systematic position within ptilodictyine cryptostomes, specifically the Family Ptilodictyidae Zittel, 1880.     

An unusual euchelicerate linking horseshoe crabs and eurypterids,from the Lower Devonian (Lochkovian) of Yunnan,China

Two new specimens of the xiphosuran Kasibelinurus yueya Lamsdell, Xue & Selden, 2013 are described, from the same horizon and locality as the type. The new specimens are conspecific with the type, but show ventral morphology, which considerably alters interpretation of the species. It can no longer be referred to Kasibelinurus Pickett, 1993, and a new genus, Houia n. gen., is erected. Houia shows a unique combination of xiphosurid and chasmataspidid/eurypterid characteristics such as lack of opisthosomal pleura and possession of a large, ventral metastomal plate. Phylogenetic analysis of all the major chelicerate orders suggests that Houia branched from the main euchelicerate lineage prior to the divergence of the three constituent clades of the Dekatriata Lamsdell, 2013 (Eurypterida, Arachnida, Chasmataspidida). Together with bunodids and pseudoniscids, Houia provides evidence for basal dekatriatans persisting into the middle Palaeozoic alongside eurypterids and arachnids and that the morphological diversity of these basal forms was greater than previously thought.     

First record of a xiphosuran trackway (<Emphasis Type="Italic">Kouphichnium</Emphasis> isp.) from the Jurassic of India

A 44 cm long trackway from the Kimmeridgian-Tithonian of the western Kachchh Basin represents the first evidence of the occurrence of members of the order Xiphosura in the Jurassic of India. The track consists of the imprints of the legs and the tail spine of a 36–44 cm long limulid and can be assigned to the ichnogenus Kouphichnium Nopsca, 1923. It was formed in a soft, silty to fine sandy substrate below the storm wave-base and preserved as a hyporelief at the base of a thin sandstone bed deposited by storm-induced currents. Eight different types of xiphosuran behaviour are known to be preserved as trace fossils: “swimming”, running, walking, crawling, mating, dying, ploughing, and resting. The present trackway from the Jurassic of India represents crawling, i.e., comparatively slow locomotion (Repichnia).     

The book lungs of Scorpiones and Tetrapulmonata (Chelicerata, Arachnida): evidence for homology and a single terrestrialisation event of a common arachnid ancestor

The question of whether Arachnida (Chelicerata) conquered terrestrial habitats only once or several times is controversial. The key group in this respect is the Scorpiones. Several authors claim that they became terrestrial independently of other arachnid lineages. This argumentation uses two lines of evidence. One is that book lungs of scorpions and other arachnids are considered non-homologous because they occur on different segments. The other line is based on fossil evidence which suggests that early scorpions were aquatic, together with a putative sister group relationship between scorpions and the aquatic Eurypterida. To address this problem we undertook a comparative scanning electron microscopical and histological study of the book lungs of scorpions, amblypygids, uropygids, and mesothelid spiders. In addition, we included the book gills of a xiphosuran. We found several detailed similarities in the book lungs shared by all arachnid taxa studied. Based on these findings we conclude that arachnid book lungs are homologous. Furthermore, we suggest that the apomorphic book lungs of arachnids indicate a single terrestrialisation event in the stem lineage leading to Arachnida.     

Genus <Emphasis Type="Italic">Eckhardites</Emphasis> Mitta (Cardioceratidae,Ammonoidea) from the Lower Callovian of the subboreal Jurassic

The genus Eckhardites Mitta 1999, with the type species Macrocephalites pavlowi Smorodina, 1928 is discussed and its diagnosis and assignment to the family Cardioceratidae are substantiated. Eckhardites is compared to the genus Macrocephalites Zittel (family Sphaeroceratidae), to which some workers presently assign the type species of Eckhardites. Three species of Eckhardites are recorded from the basal Callovian elatmae Zone of the Russian Platform, one of which (E. menzeli (Mönnig)) was originally described from the synchronous beds in Germany. A new species E. dietli sp. nov. is described.     

Campanian–early Eocene cephalopods from Kharga Oasis,Western Desert,Egypt

The Campanian–lower Eocene sedimentary succession in the Kharga Oasis yields rare cephalopods that have so far received little attention. Eight cephalopod species; six nautiloids and two ammonites, are identified in the study area. The nautiloids are referred to five genera in three families. All nautiloid species are recorded from the Paleocene and Eocene rocks, two of which are described as new, as follows: Cimomia kurkurensis nov. sp. and Deltoidonautilus hassani nov. sp. The two ammonite species are Libycoceras ismaelis (von Zittel, 1884) and Baculites ovatus Say, 1820, representing the families Sphenodiscidae and Baculitidae, respectively. Baculites anceps of Quaas (1902, non Lamarck, 1822) from the Maastrichtian of the Western Desert of Egypt is here assigned to Baculites ovatus. The palaeobiogeography of these species is studied in detail.     

Epizoan encrustation on Marsupites testudinarius – additional argument favoring an epifaunal mode of life of uintacrinoids?

Uintacrinoids (Uintacrinoidea Zittel) are among the best-known Late Cretaceous crinoids, but owing to their unusual morphology and widespread distribution their mode of life has become a subject of much discussion. Of several competing hypotheses, nektonic, pseudoplanktonic, hemipelagic, semi-infaunal and epibenthic lifestyles have been suggested. Recent study synthesizing and extending previous data has shown that these crinoids were epibenthic, holding their arms vertically for feeding. However, evidence supporting a rheophilic epibenthic model over an alternative rheophobic semi-infaunal model is still limited. Here we report epizoans, mostly represented by serpulids and bryozoans, encrusting thecal plates of Marsupites testudinarius from the Lägerdorf in Germany. Although a definitive interpretation whether recorded infestations occurred syn vivo or post mortem is not certain, it is remarkable that all epizoans (or their traces) are attached to the convex side (latera) of well-preserved isolated plates displaying no signs of reworking. Furthermore, a bryozoan colony crossing plate boundaries has been also found on the surface of a sub-articulated theca suggesting that it may have been colonised syn vivo. Recorded epibiotic associations, whether syn vivo or post-mortem, must have developed prior to burial of the specimens, above the surface of sea floor, and thus provide another argument against rheophobic semi-infaunal mode of life of uintacrinoids.     

Gigantism among Late Jurassic limulids: New ichnological evidence from the Causses Basin (Lozère,France) and comments on body-size evolution among horseshoe crabs

An abundant ichnological material composed of xiphosuran trackways and isolated traces was discovered in Upper Jurassic limestones from the Causses Basin (Causse Méjean, Lozère, France). The morphology of the imprints supports their identification as Kouphichnium isp. In contrast to the most frequent case, the trackways are composed of omnipresent pusher imprints sometime associated with leg traces, but with no telson mark. We argue that this pattern reflects actual surface traces rather than an incomplete set of undertracks. The size distribution of the sampled ichnites is broadly bimodal. This is best explained by sexual dimorphism, a phenomenon frequently observed in modern xiphosurans. Analysis of the trace fossils further suggests that several growth stages are recorded and that the horseshoe crabs were walking in a protected and flat environment like a lagoon. This area, certainly close to a mating ground, was occasionally affected by a continental influence. The biometric study of the tracks suggests a gigantic size for the trackmakers whose body may have reached 84 cm in length. This discovery complements the few reports on other gigantic horseshoe crabs in the Jurassic of Western Europe, thus casting doubt on the postulated increase in body size from the Palaeozoic to the Recent. Furthermore, a literature review shows that there are still major gaps in the record of limulid body-fossils and tracks. Thus, neither of these archives can be taken at face value for quantifying the body-size evolution of horseshoe crabs.