Morphology of Cambrian lobopodian eyes from the Chengjiang Lagerstätte and their evolutionary significance

Visual organs are widely distributed throughout the animal kingdom and exhibit a great diversity of morphologies. Compound eyes consisting of numerous visual units (ommatidia) are the oldest preserved visual systems of arthropods, but their origins are obscure and hypothetical models for their evolution have been difficult to test in the absence of unequivocal fossil evidence. Here we reveal the detailed eye structures of well-preserved Early Cambrian lobopodians Luolishania longicruris and Hallucigenia fortis from the Chengjiang Lagerstätte, China. These animals possess a pair of eyes composed of at least two visual units, interpreted as pigment cups. Contrary to previous suggestions that Cambrian lobopodians possessed ocellus-like eyes comparable to those of extant onychophorans, this multi-component structure is more similar to the lateral eyes of arthropods. Morphological comparison and phylogenetic analyses indicate that these lobopodian eyes may represent an early stage in the evolution of the ancestral visual system of euarthropods.     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

A new arthropod from the Silurian Konservat-Lagerstätte of Herefordshire, UK

A small, non-biomineralized, macrophagous arthropod with chelicerate affinities, Offacolus kingi gen. et sp. nov., from the Silurian (Wenlock Series) of Herefordshire, UK, is described. The dorsal exoskeleton comprises an arch-like cephalic shield, a thorax of three free tergites and a triangular posterior tagma of five fused tergites, the last with a stout postero-dorsally directed medial spine. Seven pairs of appendages beneath the cephalic shield surround a postero-medially sited oral cavity on the ventral surface of the head. Appendages I and, probably II are uniramous and project antero-ventrally; I was sensory and II sensory and/or ambulatory. Appendages III-VI are biramous, each with an antero-ventrally projecting ramus and a robust, highly geniculate, horizontally oriented ramus that projects through an anterior gape. The former rami were ambulatory and the latter have spinose terminal podomeres and functioned as a unit for trapping food and transferring it towards the oral cavity. Appendage VII, which is probably uniramous, is posteroventrally directed and flap like. Each tergite of the thorax and posterior tagma covers at least a pair (probably two pairs) of probably biramous appendages with each ramus flap like and setose.     

Redescription and systematic position of Diandongia pista from the Chengjiang Lagerstätte

Diandongia pista Rong is the thickest-shelled and strongly mineralized of all the brachiopods that have been recorded from the Chengjiang Fauna. It is also one of the most common species in the biota. Previous accounts have concentrated largely on the exceptionally preserved soft-bodied anatomy of this species. Here its detailed internal and external shell morphologies are described for the first time, based on BSEM and SEM observations. Its general shell morphology, pseudointerareas, and pustulose ornamentation are most similar to those of the Botsfordiidae, where many species have a Diandongia-like rhombic pattern of pustules in the juvenile apical region only. On the other hand, other aspects of the muscle pattern and the partly smooth adult ornamentation of D. pista is more suggestive of species within the Neobolidae, like Edreja. The mantle canal system in sub-adults of D. pista is comparable to that of the Eoobolidae and Obolidae, whereas the adult vascular system branches peripherally in dichotomy. D. pista may be the earliest representatives possessing some plesiomorphic characters of the Botsfordiidae.     

A soft-bodied euarthropod from the early Cambrian Xiaoshiba Lagerstätte of China supports a new clade of basal artiopodans with dorsal ecdysial sutures

We describe the exceptionally well-preserved non-trilobite artiopodan Zhiwenia coronata gen. et sp. nov. from the Cambrian Stage 3 Xiaoshiba Lagerstätte in Yunnan, China. The exoskeleton consists of a cephalic shield with dorsal sutures expressed as lateral notches that accommodate stalked lateral eyes, an elongate trunk composed of 20 tergites—the first of which is reduced—and a short tailspine with marginal spines. Appendicular data include a pair of multi-segmented antennae, and homonomous biramous trunk limbs consisting of an endopod with at least seven podomeres and a flattened exopod with lamellae. Although the presence of cephalic notches and a reduced first trunk tergite invites comparisons with the petalopleurans Xandarella, Luohiniella and Cindarella, the proportions and exoskeletal tagmosis of Zhiwenia do not closely resemble those of any major group within Trilobitomorpha. Parsimony and Bayesian phylogenetic analyses consistently support Zhiwenia as sister-taxon to the Emu Bay Shale artiopodan Australimicola spriggi, and both of them as closely related to Acanthomeridion from the Chengjiang. This new monophyletic clade, Protosutura nov., occupies a basal phylogenetic position within Artiopoda as sister-group to Trilobitomorpha and Vicissicaudata, illuminates the ancestral organization of these successful euarthropods, and leads to a re-evaluation of the evolution of ecdysial dorsal sutures within the group.     

Controls on gut phosphatisation: the trilobites from the Weeks Formation Lagerstätte (Cambrian; Utah)

Despite being internal organs, digestive structures are frequently preserved in Cambrian Lagerstätten. However, the reasons for their fossilisation and their biological implications remain to be thoroughly explored. This is particularly true with arthropods – typically the most diverse fossilised organisms in Cambrian ecosystems – where digestive structures represent an as-yet underexploited alternative to appendage morphology for inferences on their biology. Here we describe the phosphatised digestive structures of three trilobite species from the Cambrian Weeks Formation Lagerstätte (Utah). Their exquisite, three-dimensional preservation reveals unique details on trilobite internal anatomy, such as the position of the mouth and the absence of a differentiated crop. In addition, the presence of paired pygidial organs of an unknown function is reported for the first time. This exceptional material enables exploration of the relationships between gut phosphatisation and the biology of organisms. Indeed, soft-tissue preservation is unusual in these fossils as it is restricted to the digestive structures, which indicates that the gut played a central role in its own phosphatisation. We hypothesize that the gut provided a microenvironment where special conditions could develop and harboured a source of phosphorus. The fact that gut phosphatization has almost exclusively been observed in arthropods could be explained by their uncommon ability to store ions (including phosphorous) in their digestive tissues. However, in some specimens from the Weeks Formation, the phosphatisation extends to the entire digestive system, suggesting that trilobites might have had some biological particularities not observed in modern arthropods. We speculate that one of them might have been an increased capacity for ion storage in the gut tissues, related to the moulting of their heavily-mineralised carapace.     

A new look at the Late Jurassic Canjuers conservation Lagerstätte (Tithonian,Var, France)

The Canjuers conservation Lagerstätte represents a Late Jurassic lagoonal environment. The sedimentology and stratigraphy of the locality show three different depositional sequences. Fossils are mainly found in the basalmost layers that correspond to the first phase of deposition in the lithographic limestones sensu stricto. The fossil biodiversity is rich. So far, more than 1000 specimens including 38 invertebrate and 18 vertebrate taxa have been recovered from the limestones. The depositional information suggests that most invertebrates and vertebrates were not autochthonous to the lagoon, but swept in during storm events from the open sea or nearby emerged reef environments.     

Unique preservation of siliceous dinoflagellate motile cells from the Oligocene fossil Lagerstätte of Sieblos,Germany

The Triassic to Recent fossil record of the dinoflagellates is represented overwhelmingly by geologically resistant, organic-walled, non-motile resting cysts; such cysts are formed following the sexual phase in the life cycle. Very few confirmed records exist of the motile stage being preserved in the fossil record. This paper reports the occurrence of two very unusual dinoflagellate taphofacies, one developed in bituminous shales and the other in micrites, from the Oligocene fossil Lagerstätte at Sieblos, Hesse, Germany. A new dinoflagellate taxon, Sieblososphaera martini sp. nov. has been identified through analysis of dissociated skeletal elements in the bituminous shales and external moulds and casts in the micrites. The unique preservation of these fossils confirms them not only as primary biogenically silicified motile thecate cells, but also indicates that there was a much greater range of tabulation present within the subfamily Lithoperidiniaceae than has hitherto been recognized.     

A new probable stem lineage crustacean with three-dimensionally preserved soft parts from the Herefordshire (Silurian) Lagerstätte, UK

A new arthropod with three-dimensionally preserved soft parts, Tanazios dokeron, is described from the Wenlock Series (Silurian) of Herefordshire, England, UK. Serial grinding, digital photographic and computer rendering techniques yielded 'virtual fossils' in the round for study. The body tagmata of T. dokeron comprise a head shield and a long trunk. The head shield bears six pairs of horn-like spines and the head bears five pairs of appendages. The antennule, antenna and mandible are all uniramous, and the mandible includes a gnathobasic coxa. Appendages four and five are biramous and similar to those of the trunk: each comprises a limb base with an endite, an enditic membrane, and two epipodites, plus an endopod and exopod. The hypostome bears a large cone-like projection centrally, and there may be a short labrum. The trunk has some 64 segments and at least 60 appendage pairs. A very small telson has the anus sited ventrally in its posterior part and also bears a caudal furca. Comparative morphological and cladistic analyses of T. dokeron indicate a crustacean affinity, with a probable position in the eucrustacean stem group. As such the epipodites in T. dokeron are the first recorded in a eucrustacean stem taxon. The new species is interpreted as a benthic or nektobenthic scavenger.     

中国早寒武世高肌虫的最新研究--评介《A Monograph of Bradorrid Arthropods from the Lower Cambrian of Southwest China》

中国早寒武世高肌虫(Bradoriida也被翻译为金臂虫)研究已近百年历史,但最近20年来的研究成果所造成的混乱,已到了无所适从的地步.侯先光等经过5年的重新研究,其成果作为专著已在英国杂志出版发行(Hou et al.,2002).     

New sea spiders from the Jurassic La Voulte-sur-Rhône Lagerstätte

The diverse and exceptionally well-preserved pycnogonids described herein from the Middle Jurassic La Voulte Lagerstätte fill a 400 Myr gap of knowledge in the evolutionary history of this enigmatic group of marine arthropods. They reveal very close morphological and functional (locomotion, feeding) similarities with present-day pycnogonids and, by contrast, marked differences with all Palaeozoic representatives of the group. This suggests a relatively recent, possibly Mesozoic origin for at least three major extant lineages of pycnogonids (Ammotheidae, Colossendeidae, Endeidae). Combined evidence from depositional environment, faunal associates and recent analogues indicate that the La Voulte pycnogonids probably lived in the upper bathyal zone (ca 200 m). Our results point to a remarkable morphological and ecological stability of this arthropod group over at least 160 Myr and suggest that the colonization of the deep sea by pycnogonids occurred before the Jurassic.     

New Fossil Scorpion from the Chiapas Amber Lagerst?tte

A new species of scorpion is described based on a rare entire adult male preserved in a cloudy amber from Miocene rocks in the Chiapas Highlands, south of Mexico. The amber-bearing beds in Chiapas constitute a Conservation Lagerstätte with outstanding organic preservation inside plant resin. The new species is diagnosed as having putative characters that largely correspond with the genus Tityus Koch, 1836 (Scorpiones, Buthidae). Accordingly, it is now referred to as Tityus apozonalli sp. nov. Its previously unclear phylogenetic relationship among fossil taxa of the family Buthidae from both Dominican and Mexican amber is also examined herein. Preliminarily results indicate a basal condition of T. apozonalli regarding to Tityus geratus Santiago-Blay and Poinar, 1988, Tityus (Brazilotityus) hartkorni Lourenço, 2009, and Tityus azari Lourenço, 2013 from Dominican amber, as was Tityus (Brazilotityus) knodeli Lourenço, 2014 from Mexican amber. Its close relationships with extant Neotropic Tityus-like subclades such as ‘Tityus clathratus’ and the subgenus Tityus (Archaeotityus) are also discussed. This new taxon adds to the knowledge of New World scorpions from the Miocene that are rarely found trapped in amber.     

Amplectosporangium—A New Genus of Plant from the Lower Devonian of Sichuan,China

A new genus and species Amplectosporangium jiangyouense, is described from the Pingyipu Formation (Siegenian) of the Lower Devonian in Jiangyou district, Sichuan, China. The plant consists of naked dichotomizing erect axes with terminal fertile branch-systems. Sporangia are oval-shaped each with a short stalk and are arranged along the inner side of dichotomized axes in single row. By comparison with other known Devonian plants the genus is considered to represent a new morphological type of early land plants. Based on the theory of the telomic origin of the ovular integument, the plant may represent an early stage in the evolution of integument.     

Mineralization of soft-part anatomy and invading microbes in the horseshoe crab Mesolimulus from the Upper Jurassic Lagerstätte of Nusplingen, Germany

A remarkable specimen of Mesolimulus from the Upper Jurassic (Kimmeridgian) of Nusplingen, Germany, preserves the musculature of the prosoma and associated microbes in three dimensions in calcium phosphate (apatite). The musculature of Mesolimulus conforms closely to that of modern horseshoe crabs. Associated with the muscles are patches of mineralized biofilm with spiral and coccoid forms. This discovery emphasizes the potential of soft-bodied fossils as a source for increasing our knowledge of the diversity of fossil microbes in particular settings.     

The fish assemblage from the Pennsylvanian Buckhorn Asphalt Quarry Lagerstätte (Oklahoma,USA)

A diverse assemblage of fishes (isolated teeth and scales) is reported from the Middle to Upper Pennsylvanian Buckhorn Asphalt Quarry Lagerstätte in Oklahoma, USA. The assemblage includes chondrichthyans such as the bransonelliform Bransonella lingulata, the xenacanthiform Xenacanthus, the symmoriiform Stethacanthus, ctenacanthiforms, an polyacrodontid euselachian, anachronistid neoselachians (Cooleyella amazonensis and C. sp.) and an iniopterygian paraselachian cf. Sibyrhynchidae. Moreover, the assemblage encompasses remains of acanthodians and various actinopterygians and a single specimen of an osteolepiform sarcopterygian. Most of the taxa are cosmopolitan during the Late Palaeozoic. The occurrence of bransonelliform and xenacanthiform species at the same locality is very rare in the Carboniferous. The assemblage yields the first Carboniferous occurrences of a polyacrodontid hybodontoid and an actinopterygian belonging to the Acropholidae. Bransonella lingulata from the Desmoinesian/Moscovian of the Buckhorn Asphalt Quarry in Oklahoma represents the youngest occurrence of this species.     

Hsüa robusta,a new land plant from the Lower Devonian of Yunnan,China

A new psilophytic plant, Hsüa robusta, is found in the Xujiachong Formation (Emsian) of the Lower Devonian from the Qüjing (= Kütsing) district of Yunnan, China. This plant is tentatively referred to the Cooksoniaceae of Rhyniales. Hsüa gen. nov. Type species: Hsüa robusta (Li et Cai) C. S. Li. Diagnosis: Plants erect and then creeping. Main axes dividing pseudomonopodially and bearing dichotomous lateral branches which somewhat differentiate into vegeta, tire and fertile ones, with dichotomous root-like and rhizophore-like appendages. Sporangia terminal, round to reniform or wide reniform, dehiscing along distal margin into two equal halves. Spores homosporous, trilete. Stomata anomocytic. Protostele centrarch. Hsüia robusta (Li et Cai) C. S. Li, comb. nov. Cooksonia zhanyiensis Li et Cai, Acta Geologica Sinica, 52 (1) 1978, p. 10, pl. II, fig. 6.——Taeniocrada robusta Li et Cai,ib. p. 10, pl. II, fig. 7—14. Diagnosis: Characters same as in generic diagnosis. Main axes 6—10 mm wide and at least 24 cm long, with vascular strands 1.2—2.4 mm acr oss. Fertile branches 3—4 times equally or unequally dichotomous, 10—1.5 mm in width and up to 11 cm in length, possessing a vascular bundle of 0.5 mm in its greatest diameter. Branches circinately coiled in apical regions. Axial tubercles, root-like and rhizophore— like appendages arising from the main axes usually anterior to the lateral branches. Axial tubercle round with a diameter of 2.2—2.4 mm, having a vascular bundle about l mm across. Rootlike branches 3 times bifurcate, 1—0.3 mm wide and up to 1.5 cm long, with a vascular bundle about 0.1 mm across. Rhizophore-like appendages forked, 3—1.7 mm in width, possessing a vascular bundle of 0.7 mm in its greatest diameter. Root-like protuberances sometimes arising from rhilzophore-like branches. Epidermal cells of axes generally elongate, measuring 60—290μby 25—60 μ. Stomata mainly fusiform, 90—110 μ long and 50—60μ wide, consisting of a pair of guard cells enclosing a pore 6—15μ in length and 1—3μ in width. Cuticle of guard cells quite thick. Stomatal density about 5 per mm². Sporangia 0.8—4.2 mm high, 1.0—8.2 mm across, usually having a dehiscent distal border which measures 50—100μ broad. Demarcation between sporangium and its stalk quite clear. Epidermal cells of basal part of sporangial walls elongate, about 100 μ long and 30μ wide, but those of distal part isodiametrally polygonal, about 50μ in diameter. Stomata, radially arranged scattering over sporangial walls, generally round about 50μ in diameter and 50 per sporangium. Spores round, 18—36μ (average 27μ) indiameter, and smooth. Tracheids of protoxylem about 10μ across; those of metaxylem about 30μ across, with scalariform thickening. This plant is similar to Renalia hueberi Gensel in general morphology, but differs from the latter in possessing root-like and rhizophore-like branches. The generic name is derived from Prof. Hsü Jen. This paper is a thesis for M. Sc.     

Two Flat-Backed Polydesmidan Millipedes from the Miocene Chiapas-Amber Lagerst?tte,Mexico

Two species of fossil polydesmidan millipedes (Diplopoda: Polydesmida) embedded in amber are described from Miocene strata near Simojovel, in the Chiapas Highlands, Mexico. Maatidesmus paachtun gen. et sp. nov., placed into Chelodesmidae Cook, 1895, and Anbarrhacus adamantis gen. et sp. nov., assigned in the family Platyrhacidae Pocock, 1895. Morphological data from fossil specimens have been recovered using 3D X-ray micro-computed tomography and regular to infrared-reflected microscopy. Both fossil species are recognizable as new primarily but not exclusively, by collum margin modification and remarkable paranotal and metatergite dorsal sculpture.     

A new Microphorites in the Lower Cretaceous amber of the Southwest of France (Diptera: Dolichopodidae, “Microphorinae”)

A new deposit of Lower Cretaceous amber, found in Charente-Maritime (SW France) has yielded an important entomofauna with numerous arthropod associations characteristic of moist ground. We describe a new species of Dolichopodidae: ‘Microphorinae’(Diptera: Empidoidea), Microphorites deploegi n. sp. on the basis of seven male and female specimens of exceptional state of preservation. This genus was previously only known from Lebanese amber of the Lower Cretaceous. The present discovery supports a reconstruction of the palaeoenvironment as a sandy beach along the sea, under a warm climate.