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Decapod crustaceans are the most diverse group of fossil invertebrates from the Upper Cretaceous Sahel Alma Lagerstätte (Lebanon, Middle East). They are mainly represented by abundant crest−bearing shrimps which were first described as Penaeus libanensis. We review this species applying the new systematic nomenclature and we propose a more complete description based on 54 unpublished specimens. This review leads to the erection of Palaeobenthesicymus gen. nov. and to the new combination Palaeobenthesicymus libanensis that is the oldest record of the family Benthesicymidae. A neotype is herein designated. Detailed comparisons with extant analogues suggest that the crest bearing shrimps inhabited relatively deep water settings most probably exceeding 150 m, at the transition between the lower circalittoral and the upper bathyal zones, under dysphotic or aphotic conditions.
An isolated exopod in uncompressed three−dimensional “Orsten”−type preservation from the Cambrian of Australia represents a new species of Marrellomorpha, Austromarrella klausmuelleri gen. et sp. nov. The exopod is composed of at least 17 annuli. Each of the proximal annuli carries a pair of lamellae: one lamella on the lateral side and one on the median side. The distal annuli bear stout spines in the corresponding position instead of lamellae, most likely representing early ontogenetic equivalents of the lamellae. The new find extends the geographical range of the taxon Marrellomorpha. Additionally, it offers a partial view into marrellomorph ontogeny. The occurrence of a marrellomorph fragment in “Orsten”−type preservation provides new palaeo−ecological insights into the possible connections between the “Orsten” biotas and other fossil Lagerstätten. Finding such connections is necessary for understanding the complex ecosystems of early Palaeozoic times.
Three species of bradoriid arthropods from the lower to middle Cambrian transitional interval of Scania, southern Sweden, are described and illustrated: Beyrichona tinea from the top of the traditional lower Cambrian (Gislöv Formation; Ornamentaspis? linnarssoni Zone), and Hipponicharion eos and Alutella sp. from the basal portion of the traditional middle Cambrian (lowermost part of the Alum Shale Formation). The bradoriid fauna compares most closely with others previously described from western and eastern Avalonia (New Brunswick and England). The record of B. tinea suggests a correlation between the “Protolenus Zone” (Hupeolenus Zone) of western Avalonia and the O.? linnarssoni Zone of Scandinavia. Hipponicharion eos appears to be a fairly long−ranging species as it has previously been recorded from upper lower Cambrian or lower middle Cambrian strata in New Brunswick, Poland, and probably Sardinia. The record of H. eos from the lowermost part of the Alum Shale Formation suggests that this largely unfossiliferous interval in the Scanian succession is not younger than the Acadoparadoxides oelandicus Superzone. The genus Alutella has not previously been recorded from the Acado−Baltic Province.
A highly spinose fragment of a possibly raptorial appendage from the Arenig (Early Ordovician) of the Upper Fezouata Formation north of Zagora, southeastern Morocco is described as the arthropod Pseudoangustidontus duplospineus gen. et sp. nov. The single fragmentary specimen displays a unique morphology, carrying at least 39 pairs of spines (i.e., 78 spines) of very regularly alternating lengths. Pseudoangustidontus gen. nov. shows some similarities to a number of spinose arthropod appendages and appendage parts, most notably to the spine−bearing podomeres of the third prosomal appendage of megalograptid eurypterids and the problematic and incompletely known genus Angustidontus. However, megalograptids and Angustidontus both have a lower spine count, while the latter also carries only a single row of spines. Because no known arthropod displays a morphology closely comparable to that of Pseudoangustidontus gen. nov., the affinities of the new fossil within Arthropoda remain uncertain.
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Zhenghecaris shankouensis gen. et sp. nov. is one of the largest “bivalved” arthropods of the Lower Cambrian Maotianshan Shale fauna. Its non−mineralized carapace was dome−like, laterally compressed, armed with rostral features, and probably enclosed the entire body of the animal. Zhenghecaris was provided with elliptical stalked lateral eyes. The carapace design, external ornament and visual organs of Zhenghecaris suggest affinities with the Thylacocephala, an extinct (Lower Silurian to Upper Cretaceous) group of enigmatic arthropods whose origins remain poorly understood. The bivalved arthropodsIsoxys and Tuzoia (Lower and Middle Cambrian) are two other potential thylacocephalan candidates making this group of arthropods a possible new component of Cambrian marine communities. Zhenghecaris, Isoxys, and Tuzoia are interpreted as nektonic animals that probably inhabited the lower level of the water column in shallow shelf settings at depths of perhaps 100–150 m or less. Their feeding mode either in the water column (e.g., mesozooplankton) or on the substrate (e.g., small epibenthos, detritus) is uncertain, although some of these arthropods were possibly mid−water predators (e.g., Isoxys with raptorial appendages).
Onychodictyon ferox, from the Lower Cambrian Chengjiang Lagerstätte, is a worm−like armored lobopod with 10 pairs of dorsal plates and a pair of probable sensory appendages (“antennae”). Newly discovered complete specimens show that Onychodictyon is represented by two species: O. ferox with a “head” bearing a pair of dorsal “antennae” and a trunk with 11 pairs of limbs, whereas O. gracilis has a blunt anterior end without any appendages and a trunk with 12 pairs of limbs. Because of the close resemblance of the “antennae” of O. ferox and those of the lobopod Miraluolishania, they are considered to be homologous structures. The “antennae” of Cambrian lobopodians are proposed to be homologous with the arthropod antennulae.
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Open antra, a special form of external brood pouches in Ordovician ostracodes suitable for both egg and brood care, are enabled by the associated pore system. Special kinds of radial pores connected with the adventral sculptures are described for the first time. Relationships exist between antral development and lifestyle in that open antra occur in nektobenthic taxa, while closed antra (false brood pouches) are connected with a benthic lifestyle. Taxa, particularly those with open antra, which are morphologically very similar in being non−lobate or non−sulcate, can be distinguished by the construction of the respective antra and the associated pore systems, as exemplified by Levisulculus, Swantina, and Ampletochitina.
The La Voulte Lagerstätte is remarkable for its unique soft−bodied fauna (e.g., worms, coleoid squids) and its exceptionally preserved arthropods mainly found in small sideritic concretions. This arthropod fauna includes 30 different species assigned to the crustaceans, the thylacocephalans and the pycnogonids. Crustaceans are the most diversified group with 23 species distributed in a dozen families. Quantitative analyses based on 388 nodules reveals four dominant groups: (i) the enigmatic thylacocephalan arthropods (33%), (ii) the Solenoceridae shrimps (22%), (iii) the Coleiidae crustaceans (15%), and (iv) the Penaeidae shrimps (10%). Converging lines of evidence from depositional environment and modern analogues, indicate that this arthropod fauna probably inhabited a deep water setting most probably exceeding 200 m (= bathyal zone) under dysphotic or aphotic conditions. This new set of data sheds new light on the deep−sea colonisation by animal communities in the Mesozoic.
The enigmatic marrellomorph arthropod Furca bohemica from the Upper Ordovician Letná Formation, is redescribed. Based on existing museum specimens and new material collected from the southern slope of Ostrý Hill (Beroun, Czech Republic), the morphology and taphonomy of F. bohemica is reappraised and expanded to produce a new anatomical interpretation. The previously distinct taxa F. pilosa and Furca sp., are synonymised with F. bohemica, the latter being represented by a tapho−series in which decay has obscured some of the diagnostic features. A cladistic analysis indicates close affinities between F. bohemica and the Hunsrück Slate marrellomorph Mimetaster hexagonalis, together forming the Family Mimetasteridae, contrary to previous models for marrellomorph internal relationships. As with other representatives of the group, the overall anatomy of F. bohemica is consistent with a benthic, or possibly nektobenthic, mode of life. The depositional setting of the Letná Formation indicates that F. bohemica inhabited a shallow marine environment, distinguishing it palaeoecologically from all other known marrellomorphs, which have been reported from the continental shelf.
The new fossil thrips Uzelothrips eocenicus P. Nel and A. Nel sp. nov. (Thysanoptera: Uzelothripidae) is described from two lowermost Eocene amber−preserved specimens (one macropterous and one apterous). The family Uzelothripidae is only known so far from a single extant species, Uzelothrips scabrosus. The fossils differ from the extant species only by the antennal segments III and IV, which appear distinctly separated instead of being fused as in the in the extant U. scabrosus. Dark−coloured hyphae and conidia of the Dothideomycetes (Ascomycota) which are likely to belong to the sooty moulds (Capnodiales) are attached to the apterous fossil specimen. We consider this arthropod−fungus association not to be accidental since these fungi are also found in extant specimens of these uzelothripids, suggesting very specific long−term interactions and strong habitat specificity.
A new myodocope ostracod Sineruga insolita gen. et sp. nov. is herein described from the Armorican Massif (France). Sineruga resembles Silurian myodocopes (i.e., bolbozoids and cypridinids) in having anterior features (thinner carapace) possibly related to vision. On the other hand, it resembles entomozoid ostracods in having a bean−shaped outline along with a deep adductorial sulcus and a simple muscle spot, but lacks their characteristic ribbed ornament. The data available suggest that Sineruga insolita was probably an atypical, non−ribbed member of the entomozoids thus indicating that early entomozoids may have had smooth representatives. Comparisons with other Recent and fossils ostracods show that the presence of a rostrum and/or symmetrical vision related carapace features (i.e., indicating lateral eyes) can be used as diagnostic characters for myodocope ostracods. The position and shape of the dorsal connection and to a lesser extent that of the sulcus and the adductorial muscle scar can be used to discriminate the higher groups of Silurian myodocopes (bolbozoids, cypridinids, and entomozoids).
The Leanchoiliidae is well−known from abundant material of Leanchoilia, from the Burgess Shale and Chengjiang Konservat−Lagerstätten. The first Australian member of the group is Oestokerkus megacholix gen. et sp. nov., described from the Emu Bay Shale (Cambrian Series 2, Stage 4), at Buck Quarry, Kangaroo Island, South Australia, and is intermediate in age between the well known leanchoiliid species from the Burgess Shale and Chengjiang. Phylogenetic analysis of “short great appendage” arthropods (Megacheira) in the context of the chelicerate stem group resolves the Australian species as sister to Burgess Shale, Utah, and Chengjiang Leanchoilia species, but most readily distinguished from Leanchoilia and Alalcomenaeus by a different telson shape, interpreted as being forked, widening distally, and with a few dorsally curved spines at the posterior angle. Leanchoiliid interrelationships are stable to alternative character weights, and Megacheira corresponds to a clade in most analyses.
More than forty specimens from the middle Cambrian Burgess Shale reveal the detailed anatomy of Isoxys, a worldwide distributed bivalved arthropod represented here by two species, namely Isoxys acutangulus and Isoxys longissimus. I. acutangulus had a non−mineralized headshield with lateral pleural folds (= “valves” of previous authors) that covered the animal’s body almost entirely, large frontal spherical eyes and a pair of uniramous prehensile appendages bearing stout spiny outgrowths along their anterior margins. The 13 following appendages had a uniform biramous design—i.e., a short endopod and a paddle−like exopod fringed with marginal setae with a probable natatory function. The trunk ended with a flap−like telson that protruded beyond the posterior margin of the headshield. The gut of I. acutangulus was tube−like, running from mouth to telson, and was flanked with numerous 3D−preserved bulbous, paired features interpreted as digestive glands. The appendage design of I. acutangulus indicates that the animal was a swimmer and a visual predator living off−bottom. The general anatomy of Isoxys longissimus was similar to that of I. acutangulus although less information is available on the exact shape of its appendages and visual organs. I. longissimus is characterized by extremely long anterior and posterior spines. There are now seven Isoxys species known with soft−part preservation, I. acutangulus, I. longissimus from the Burgess Shale, I. auritus and I. curvirostratus from the Maotianshan Shale of China, I. communis and I. glaessneri from the Emu Bay Shale of Australia and I. volucrisfrom Sirius Passet in Greenland. The frontal appendages of Isoxys strongly resemble those of other Cambrian arthropods, characterized by a single pair of “great appendages” with a shared prehensile function yet some variability in length and shape.
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The synziphosurine horseshoe crab Pasternakevia was until now known only from a single specimen. Herein we describe three new specimens from the outcrop close to the type locality of monotypic P. podolica that show several features that were unknown or only assumed in this genus. These characters include: a smooth surface of the carapace; an opisthosoma composed of ten segments, the first one being a microtergite (strongly reduced first opisthosomal segment usually hidden under the carapace), two or three last segments fused into a metasoma, and the telson. Pasternakevia is included in Bunodidae due to a hypertrophied tergite 2, but it is linked to the Pseudoniscidae by the shape of the opisthosoma.
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