Variable preservation potential and richness in the fossil record of vertebrates

Variation in preservation and sampling probability clouds our estimates of past biodiversity. The most extreme examples are Lagerstätten faunas and floras. Although such deposits provide a wealth of information and represent true richness better than other deposits, they can create misleading diversity peaks because of their species richness. Here, we investigate how Lagerstätten formations add to time series of vertebrate richness in the UK, Germany and China. The first two nations are associated with well-studied fossil records and the last is a country where palaeontology has a much shorter history; all three nations include noted Lagerstätten in their fossil records. Lagerstätten provide a larger proportion of China's sampled richness than in Germany or the UK, despite comprising a smaller proportion of its fossiliferous deposits. The proportions of taxa that are unique to Lagerstätten vary through time and between countries. Further, in all regions, we find little overlap between the taxa occurring in Lagerstätten and in ‘ordinary’ formations within the same time bin, indicating that Lagerstätten preserve unusual faunas. As expected, fragile taxa make up a greater proportion of richness in Lagerstätten than the remainder of the fossil record. Surprisingly, we find that Lagerstätten account for a minority of peaks in the palaeodiversity curves of all vertebrates (18% in the UK; 36% in Germany and China), and Lagerstätten count is generally not a good overall predictor of the palaeodiversity signal. Vastly different sampling probabilities through taxa, locations and time require serious consideration when analysing palaeodiversity curves.     

Preservational bias controls the fossil record of pterosaurs

Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeo‐diversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group's evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non‐significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large‐bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic–Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution.     

The La Voulte-sur-Rhône Lagerstätte (Middle Jurassic,France)

The La Voulte Lagerstätte (Jurassic, France) is characterized by a diverse and exceptionally well-preserved fauna. It was located along the western Tethyan margin where the submarine relief was steep. The sedimentation was dominated by marls suggesting a low energy and deep-water depositional environment. The La Voulte biota is remarkable for its biodiversity with about 60 different species. The major components are the arthropods (50% of the species), the cephalopods (10%) and the echinoderms (10%). Among the arthropods, the composition and some anatomical features suggest a relatively deep-water environment characterized by dim-light conditions. The cephalopods and other organisms (pycnogonids, asterids), have extant analogues that all live in deep-water niches always exceeding 200 m. The fauna as a whole bears the characteristic features of the present-day bathyal fauna. The La Voulte Lagerstätte may be one of the rare Jurassic Lagerstätten, if not the unique, to have fossilized a deep marine fauna.     

Diversity and palaeoecology of a highly diverse Late Triassic marine biota from the Cassian Formation of north Italy

A sample of marine invertebrates from the Late Triassic Cassian Formation (north Italy) yielded one of the most diverse Early Mesozoic fossil assemblages ever reported (c. 170 species). The assemblage was found in basin clays, but was transported from nearby carbonate platforms as indicated by fragmentation, microbial encrustation and the presence of coated grains and ooids. Most of the specimens are small (< 1 cm) reflecting both, small adult sizes and size sorting during transport. Rarefaction analysis suggests that diversity of surface collection and bulk sampling is the same. However, rank abundance, species richness and taxonomic composition differ strongly according to sampling method. Low‐grade lithification of the sediments is the main reason that high diversity can be recognized, because it facilitates disaggregation and finding of small molluscs. Sample standardization shows that the studied assemblage is much more diverse than known Early Triassic assemblages. However, its diversity is similar to that of Anisian assemblages. This suggests that recovery from the end‐Permian mass‐extinction was quite advanced in the Middle Triassic and alpha‐diversity remained high until the Late Triassic. According to current models, Early Triassic and Anisian faunas match the niche overlap phase of recovery during which diversity is built up by increasing alpha‐diversity, whereas beta‐diversity rises slowly. Subsequently, habitat width of species contracts because of increasing competition, making beta‐diversity the principal drive of overall diversity increase. The diversity pattern of various Late Triassic Cassian associations meets the predictions for the transition from the niche overlap to the habitat contraction phase.: Triassic, Cassian Formation, palaeoecology, diversity, mollusc dominance.     

Facies of two important Early Triassic gastropod lagerstätten: implications for diversity patterns in the aftermath of the end-Permian mass extinction

Two important lagerstätten of Early Triassic gastropods, the Sinbad Limestone (Utah, USA) and the Gastropod Oolite (North Italy) yield about 40% of all described Early Triassic species. This great contribution to the global diversity and the exceptional good preservation render high information content, which characterizes fossil lagerstätten. The Smithian Sinbad Limestone contains the most diverse Early Triassic gastropod fauna. At the type locality, it occurs in single, probably storm-induced shell bed within a series of high energy deposits underlain by intertidal microbial mats and subtidal oolite/peloid shoals. The main shell bed contains about 40 invertebrate taxa. Gastropods, scaphopods, and bivalves are most abundant and form an assemblage, which is dominated by small neritaemorphs, the opisthobranch Cylindrobullina convexa and the scaphopod Plagioglypta (annulated tubes). This assemblage lived on shallow, subtidal soft-bottoms based on sedimentological and ecological characteristics. The Dienerian (to Smithian?) Gastropod Oolite Member (North Italy) has extremely abundant, probably salinity-controlled gastropod faunas with low species richness. Almost monospecific assemblages of Pseudomurchisonia kokeni as well as assemblages with about four species are present in the Gastropod Oolite. Modern hydrobiid mudsnail faunas which are adapted to strongly fluctuating salinity in intertidal to shallow subtidal coastal areas form probably a suitable model for the Gastropod Oolite biota. Gastropods from the Werfen- and Moenkopi-Formation lagerstätten are well preserved compared to other Early Triassic deposits. The high contribution to the global diversity of just two sites suggests very incomplete sampling and preservational bias. However, the low richness of the major faunas reflects depauperate Early Triassic faunas and slow recovery from the Permian/Triassic crisis.     

Skeletal completeness of the non‐avian theropod dinosaur fossil record

Non‐avian theropods were a highly successful clade of bipedal, predominantly carnivorous, dinosaurs. Their diversity and macroevolutionary patterns have been the subject of many studies. Changes in fossil specimen completeness through time and space can bias our understanding of macroevolution. Here, we quantify the completeness of 455 non‐avian theropod species using the skeletal completeness metric (SCM), which calculates the proportion of a complete skeleton preserved for a specimen. Temporal patterns of theropod skeletal completeness show peaks in the Carnian, Oxfordian–Kimmeridgian and Barremian–Aptian, and lows in the Berriasian and Hauterivian. Lagerstätten primarily drive the peaks in completeness and observed taxonomic diversity in the Oxfordian–Kimmeridgian and the Barremian–Aptian. Theropods have a significantly lower distribution of completeness scores than contemporary sauropodomorph dinosaurs but change in completeness through time for the two groups shows a significant correlation when conservation Lagerstätten are excluded, possibly indicating that both records are primarily driven by geology and sampling availability. Our results reveal relatively weak temporal sampling biases acting on the theropod record but relatively strong spatial and environmental biases. Asia has a significantly more complete record than any other continent, the mid northern latitudes have the highest abundance of finds, and most complete theropod skeletons come from lacustrine and aeolian environments. We suggest that these patterns result from historical research focus, modern climate dynamics, and depositional transportation energy plus association with conservation Lagerstätten, respectively. Furthermore, we find possible ecological biases acting on different theropod subgroups, but body size does not influence theropod completeness on a global scale.     

A new rhynchonelliform brachiopod Longtancunella with soft-part preservation from the Hongjingshao Formation (Cambrian Stage 3) in Yunnan,South China

Rhynchonelliform brachiopods made their first appearance in early Cambrian, and became a major group within the palaeozoic evolutionary fauna since late Cambrian. Exceptionally preserved fossils from the early Cambrian Lagerstätten provide valuable chances to investigate their phylogeny and ecology. Longtancunella is one of the most interesting early rhynchonelliforms, and has been mainly recovered from the Chengjiang Lagerstätte (Series 2, Stage 3). Here, we report a new rhynchonelliform Longtancunella xiazhuangensis n. sp. from the lower Hongjingshao Formation (upper Stage 3) in Yunnan Province, China. These specimens were well preserved with soft parts, including pinnate mantle canal system and a pedicle. It is identified as a new species based mainly on its difference in shell ornamentations, pinnate mantle canals and pedicle morphology from the type species. Its pedicle looks unusually stout with distinct annulated lamellae on the surface, and reveals crucial evidence in illustrating its ecology and settling strategy as an early marine epifauna. The ecological interaction between L. xiazhuangensis and other marine animals also provides insights into the food web structure in the early Cambrian.     

Quantifying the completeness of the bat fossil record

Bats (Chiroptera) are one of the most successful extant mammalian orders, uniquely capable of powered flight and laryngeal echolocation. The timing and evidence for evolution of their novel adaptations have been difficult to ascertain from the fossil record due to chronological gaps and the fragmentary nature of most fossil bat material. Here, we quantify the quality of the bat fossil record using skeletal and character completeness metrics, which respectively document for each taxon what proportion of a complete skeleton is preserved, and the proportion of phylogenetic characters that can be scored. Completeness scores were collected for 441 valid fossil bat species in 167 genera from the Eocene to the Pleistocene. All metrics record similar temporal patterns: peak completeness in the Lutetian stage reflects the presence of Lagerstätten, while subsequent stages have very low completeness, except an Aquitanian high and a Pleistocene peak in skeletal completeness. Bat completeness is not correlated with intensity of sampling through geological time but has a weak negative correlation with publication date. There is no correlation between taxonomic richness and completeness, as the bat record predominately consists of diagnostic but isolated teeth. Consequently, bat skeletal completeness is the lowest of any previously assessed tetrapod group, but character completeness is similar to parareptiles and birds. Bats have significantly higher character completeness in the northern hemisphere, probably due to heightened historical interest and presence of Lagerstätten. Taxa derived from caves are more complete than those from fluviolacustrine and marine deposits, but do not preserve highly complete specimens.     

New trace fossil evidence for eurypterid swimming behaviour

We describe a recently discovered trace fossil from a eurypterid Konservat‐Lagerstätte in the upper Silurian Tonoloway Formation of Pennsylvania, and formally describe contemporaneous traces from the Williamsville Formation Lagerstätte of Ontario. The traces from both localities are assigned here to Arcuites bertiensis igen. et isp. nov. Based on comparisons with previously described eurypterid trackways, neoichnological experiments, and the co‐occurrence with eurypterid remains, Arcuites is interpreted as having been made by the swimming leg (sixth prosomal appendage) of swimming juvenile to adult eurypteroid eurypterids, and represents the first unambiguous trace fossil evidence for eurypterid swimming behaviour. The morphology of Arcuites indicates that eurypteroid eurypterids swam using drag‐based rowing, whereby the animal propelled itself forward by moving its oar blade‐like swimming paddles in an in‐phase backstroke. Arcuites morphology also indicates that the eurypteroid swimming appendage had a greater degree of movement than was previously suggested, and a revised rowing model is proposed. Differences in the abundance of A. bertiensis in the Tonoloway and Williamsville formations suggest a bathymetric control on eurypterid swimming behaviour and trace production. The association of Arcuites with eurypterid body fossils in both units indicates that these Lagerstätten were autochthonous assemblages and provides additional evidence for eurypterid inhabitation of shallow subtidal marine environments in the late Silurian.     

Buoyancy mechanisms limit preservation of coleoid cephalopod soft tissues in Mesozoic Lagerstätten

Coleoid cephalopods are characterized by internalization of their shell, and are divided into the ten‐armed Decabrachia (squids and cuttlefish) and the eight‐armed Vampyropoda (octopuses and vampire squid). They have a rich fossil record predominantly of the limited biomineralized skeletal elements they possess: arm hooks, statoliths, mouthparts (the buccal mass) and internal shell (gladius or pen), although exquisitely preserved soft tissue coleoids are known from several Lagerstätten worldwide. Recent studies have shown that although morphological similarities between extant decabrachian gladii and fossil examples exist, no known examples of fossil decabrachians are currently known. However, molecular clock data and phylogenetic bracketing suggest that they should be present in Lagerstätten that are rich in vampyropod soft tissue fossils (i.e. Hâkel and Hâdjoula Lagerstätten, Cretaceous, Lebanon). We propose that a hitherto unknown taphonomic bias pertaining to the differing methods of buoyancy control within coleoid groups limits preservation potential. Both negatively and neutrally buoyant decabrachians use chemical buoyancy control (ammonia) whereas vampyropods do not. In the event of rapid burial in an environment conducive to exceptional preservation, ammonia dramatically decreases the ability of the decabrachian carcass to generate the required pH for authigenic calcium phosphate replacement, limiting its preservation potential. Moreover, the greater surface area and comparatively fragile dermis further decrease the potential for fossilization. This taphonomic bias may have contributed to the lack of preserved labile soft‐tissues in other cephalopods groups such as the ammonoids.     

A new early Cambrian bivalved euarthropod from Yunnan,China and general interspecific morphological and size variations in Cambrian hymenocarines

The hymenocarines were a major and diverse group of euarthropods featured by a bivalved head carapace and a segmented trunk comprising tergo-pleural rings notably from the Cambrian Burgess Shale-type Lagerstätten. Here we document a new ~50 mm long hymenocarine Xiazhuangocaris chenggongensis n. gen. n. sp. from the Xiazhuang fossil assemblage, a slightly younger occurrence of the Chengjiang Lagerstätte within the lower Cambrian Hongjingshao Formation (Cambrian Series 2, Stage 3) in Chenggong, Yunnan, southwestern China. X. chenggongensis is characterized by the bivalved carapace with each valve bearing a protruded anterior tip and a prominent posterior notch, which resembles the P-element of the radiodont Hurdia and is unique among the hymenocarines. The trunk consists of at least 13 tergo-pleural rings including presumably eight abdominal ones terminating in a pair of ovate caudal rami. With the body organization and morphological features closest to waptiids, X. chenggongensis represents a ‘larger’ waptiid-like hymenocarine. The morphometric analysis recognizes the family-level groupings of Cambrian hymenocarines based on the number of tergo-pleural rings and the proportion of exposed trunk length, indicating that these are useful characters for studying hymenocarine morphology and phylogeny. Cambrian hymenocarines exhibited great interspecific body size variations even between sister species of the same family, further suggesting that niche diversification happened on the family level and even among phylogenetically close genera or species during the Cambrian explosion.     

Experimental investigation of insect deposition in lentic environments and implications for formation of Konservat Lagerstätten

Terrestrial insects are often remarkably well preserved in lacustrine Konservat Lagerstätten. However, the assumption that carcasses should sink fast through the water column seems contradictory as this scenario is unlikely due to excessive buoyancy and surface tension. The mechanisms that promote rapid and permanent emplacement onto the sediment surface (RPESS) of such terrestrial animal remains are not fully understood. Here we use taphonomic experiments to show that floating in water, growth of microbial biofilms and reception of rapid sediment load promote RPESS of terrestrial insect remains in lentic water bodies. Our results show that the optimum conditions for RPESS occur when terrestrial insects enter a lentic water body in articulation, experience brief decay in association with growth of microbes, then are buried rapidly by airborne volcanic ash. These results provide a model for preservation of articulated terrestrial insects and emphasize the importance of microbial activity and volcanism for insect preservation in lacustrine Konservat Lagerstätten.     

Phylogenetic structure of the extinction and biotic factors explaining differential survival of terrestrial notosuchians at the Cretaceous–Palaeogene crisis

Although the clade Crocodylomorpha is represented by few extant species (Crocodylia), it has a rich fossil record. Hundreds of species, adapted to terrestrial, semi-aquatic and marine environments, have existed over more than 200 million years. Numerous studies have attempted to characterize the factors driving the diversification and extinction events of Crocodylomorpha, resulting in ambiguous and even contradictory conclusions, which points to the need for phylogenetically and temporally smaller-scaled studies. Here, we focus on differential survival at the Cretaceous–Palaeogene (K–Pg) crisis of Notosuchia, a diverse clade of mostly terrestrial Crocodylomorpha that achieved great diversity during the Cretaceous. More precisely, we tested the effect of body size and palaeotemperatures on notosuchian survival probability during the K–Pg crisis as well as the effect of diet on the evolution of their body size. We find that Notosuchia showed an evolutionary trend towards larger body sizes through time, associated with a shift from an omnivorous to a carnivorous diet. This may explain why sebecids were the only notosuchians to survive the K–Pg crisis. We also corroborate the conclusions of previous studies that detected a Lagerstätten effect occurring in the Adamantina Formation (Upper Cretaceous, Brazil, Bauru Group). This work confirms the value of more finely-scaled macroevolutionary studies for understanding the history of a rich and complex group such as Crocodylomorpha.     

Diversity,palaeoecology and systematics of a marine fossil assemblage from the Late Triassic Cassian Formation at Settsass Scharte,N Italy

A marine fossil assemblage from the Late Triassic (Early Carnian) Cassian Formation is reported. It was retrieved by bulk sampling, including wet sieving and quantitative picking, and by quantitative surface collection. The collection consists of c. 460 specimens (foraminiferans not included) representing 54 species. In terms of abundance and species richness, it is strongly dominated by molluscs, especially gastropods. 97 % of the individuals are molluscs. The most abundant species are the gastropods Goniospira armata, Schartia carinata and Helenostylina convexa, followed by the scaphopod Plagioglypta undulata and the bivalve Palaeonucula strigilata. Disarticulated echinoderm ossicles (mostly echinoids, crinoids, few holothurians) comprise almost all of the rest of the assemblage. The studied assemblage shows moderate diversity, similar to those of previously reported assemblages or associations from basinal settings. However, it differs considerably in taxonomic composition from previously described associations of the Cassian Formation. The abundance of small gastropods is a result of their primary abundance in these ancient living communities and of the sampling method (sieving at 0.5 mm), because most of the previous associations were obtained by performing surface collections, in which small gastropods are easily overlooked. The fauna is interpreted as an autochthonous soft bottom assemblage dominated by species that lived in low epifaunal to shallow infaunal habitats. Detritivory, deposit feeding and microcarnivory represent the main feeding types. Most of the species were fully motile but slow, and either infaunal (scaphopods, nuculids, the gastropod Domerionina) or epifaunal (most other gastropods, echinoids). The present assemblage underlines the pronounced heterogeneity of the Cassian biota. The low grade of lithification and diagenetic alteration facilitates bulk sampling and the investigation of small species. This minimizes possible sampling and preservation biases, so the studied assemblage reflects the alpha diversity of this ancient living community to an unusually high degree. The following gastropod taxa are new: Helenostylina convexa n. sp., Schartia carinata n. gen. n. sp., Schartiinae n. subfam.; Cassianastraea Bandel non Volz is replaced with Bandelastraea nom. nov.     

Spinicaudatans and ostracods (Crustacea) from the Montceau Lagerstätte (Late Carboniferous, France): morphology and palaeoenvironmental significance

Spinicaudatans and ostracods form two components of the diverse arthropod fauna from the Montceau Lagerstätte (Stephanian, France). Spinicaudatans are represented by Montcestheria orri gen. and sp. nov. and Euestheria feysi sp. nov., and ostracods by a single species, Carbonita sp. aff. salteriana (Jones, 1862). Allied forms such as Montcestheria sp. aff. orri, Montcestheria sp. and Euestheria cebennensis (Grand'Eury, 1890), all from coeval localities in France, are also described. Montcestheria gen. nov. has carapace features, external (possibly sexual) dimorphism, preserved soft parts (e.g. appendages, gut) and resting eggs similar to those of Recent spinicaudatans, suggesting comparable lifestyles, reproductive strategies and feeding modes. Detailed anatomical comparisons are made with Cyzicus tetracerus from Recent ephemeral freshwater ponds. The ostracods belong to the Superfamily Carbonitoidea, which is a recurrent component of Carboniferous non‐marine biotas. Spinicaudatan‐rich assemblages typically occur in several Westphalian–Stephanian Lagerstätten (including Montceau) and localities from Europe and North America, where freshwater conditions prevailed, indicating that the group had already colonized continental waters by the Late Carboniferous. Similarities with the fauna from Recent temporary freshwater ponds (e.g. low diversity/high density spinicaudatan‐ostracod populations, synchronous spinicaudatan populations developing from resting eggs, high diversity/low density insects, amphibians) suggest the presence of temporary or ephemeral aquatic environments at Montceau as part of a complex limnic ecosystem. Flooding may have been the main driving force by which faunal and floral elements drifted away from their respective biotopes into the depositional areas, thus explaining the co‐occurrence of terrestrial (e.g. myriapods, scorpions, plants), amphibian and aquatic (e.g. conchostracans and syncarids from temporary and permanent settings, respectively) elements in fossil assemblages.     

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

A synoptic review of the Eocene (Ypresian) cartilaginous fishes (Chondrichthyes: Holocephali,Elasmobranchii) of the Bolca Konservat-Lagerstätte,Italy

Here, we review and discuss the records and taxonomy of the Ypresian (Eocene) chondrichthyans from the famous Bolca Konservat-Lagerstätte in northeastern Italy. Despite the outstanding diversity and the numerous studies focusing on the actinopterygian faunas from Pesciara and Monte Postale, the current knowledge about the systematics, taxonomy and phylogenetic relationships of the cartilaginous fishes from these Eocene sites remains elusive and largely inadequate. The celebrated Eocene Bolca Lagerstätte has yielded several exquisitely preserved articulated remains of chondrichthyan fishes in which delicate structures and soft tissues are preserved, as well as isolated teeth. The cartilaginous fish assemblage of Bolca comprises at least 17 species-level taxa belonging to 10 families in 6 orders, including selachians (Carcharhiniformes, Lamniformes), batoids (Torpediniformes, Myliobatiformes, Rajiformes) and holocephalans (Chimaeriformes). The occurrence of holocephalans represented by an isolated fin-spine of the chimeroid Ischyodus in the Bolca assemblage is reported here for the first time and represents the first record of chimeroids in the Eocene of Italy and also southern Europe. The Bolca chondrichthyan assemblage is remarkably different from those of other contemporaneous Boreal or Tethyan deposits, suggesting that its taxonomic composition is largely influenced by the palaeoenvironmental context. However, this synoptic review also highlights the importance of detailed revisions of all chondrichthyan remains from the Bolca Konservat-Lagerstätten.     

Fossil-Lagerstätten,palaeoecology and preservation of invertebrates and vertebrates from the Devonian in the eastern Anti-Atlas,Morocco

In some Devonian strata in the eastern Anti-Atlas, fossil invertebrates are abundant, display a high taxonomic diversity and indicate many shifts in palaeoecology. This is reflected in changes in faunal composition of invertebrates and vertebrates. Fossils of jawed vertebrates of late Lochkovian and younger age have been recorded and are relatively common with their abundance and diversity increasing towards the Late Devonian. Environmental changes in the Devonian also left their mark in the preservation of vertebrates and invertebrates from the Anti-Atlas, which varies strongly through time and regionally. This variation partially reflects environmental changes linked with the evolution of small marine basins during the disintegration of the continental shelf of Gondwana in this region, fluctuations of the regional sea level and other environmental changes. To improve our understanding of these ecological changes, of shifts in preservation through the succession and of the formation of Fossil-Lagerstätten, we analysed the mineral composition of some invertebrate and vertebrate samples of Devonian and Early Carboniferous age by Raman spectroscopy and X-ray diffraction. Additionally, we characterized some of these Fossil-Lagerstätten using palaeontological and sedimentological parameters. We examined eight Devonian Konzentrat-Lagerstätten and two Konservat-Lagerstätten with soft-tissue preservation (the Famennian Thylacocephalan Layer and the Hangenberg Black Shale of the southern Maïder). The last two are the first Konservat-Lagerstätten described from the Devonian of North Africa. The taphonomic and oceanic settings suggest that these Konservat-Lagerstätten are formed because of stagnation (related to vertical restriction of water exchange and water depth rather than limited spatial water exchange and a lateral restriction) in the relatively small Maïder Basin with limited water exchange with the neighbouring Tafilalt Basin. The temporally low oxygen levels in the Maïder Basin are a possible reason for the reduced chondrichthyan diversity (missing demersal and shallow water species) compared to the Tafilalt Platform.     

A cryptic record of Burgess Shale‐type diversity from the early Cambrian of Baltica

Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).