Larval and juvenile gastropods from a Carboniferous black shale: palaeoecology and implications for the evolution of the Gastropoda

A horizon in the late Visean Ruddle Shale from Arkansas contains the oldest well-preserved gastropod protoconchs known from the Americas. The gastropod fauna consists of a diverse larval shell assemblage and a low diversity assemblage of juvenile gastropods that probably had a benthic life habit. Gastropod larval shells are always isolated, i.e. the gastropods did not complete their life cycle (no metamorphosis) and were unable to become benthic. This was caused by unfavorable environmental conditions on the soft muddy bottom that was probably due to anaerobic to exaerobic conditions. The absence or scarcity of bioturbation caused by invertebrate detritus or sediment feeders in both shale and concretions (formed before compaction) favored preservation of the delicate larval shells. The lack or scarcity of infauna and bioturbation as well as the low diversity of the presumed benthos supports an interpretation of a quasi-anaerobic to exaerobic benthic environment. The superbly preserved larval shells demonstrate that there are more caenogastropod clades present in the late Palaeozoic than suggested previously. Some larval shell types have an openly coiled first whorl followed by a planktotrophic larval shell; openly coiled initial whorls are unknown from modern caenogastropods. The vetigastropods have a smooth protoconch of two whorls clearly demarked from the following whorls - a pattern unknown in modern vetigastropods which have a protoconch of less than one whorl and build no larval shell during their planktonic stage. This could indicate a link between Palaeozoic vetigastropods and the caenogastropods.     

An Evaluation of the Recently Proposed Palaeozoic Gastropod Subclass Euomphalomorpha

The recently proposed subclass Euomphalomorpha Bandel and Frýda, 1998 was primarily based on an openly coiled, planispiral initial whorl. Here, the protoconch of Carboniferous euomphalids is described and figured in detail previously unknown. This protoconch comprises distinctly less than one planispiral whorl with an abrupt transition to the telecoconch. It is umbilicate or openly coiled. Umbilicate or bilaterally symmetrical protoconchs are still present in Recent species of the Cocculiniformia, the Neomphalidae and the Docoglossa. A previously suggested close phylogenetic relationship between the euomphalids and primitive gastropod limpets is therefore corroborated by the protoconch morphology. The fact that several Palaeozoic gastropods, unlike most modern gastropods, have an openly coiled initial whorl is highly remarkable, but the meaning of this feature for the phylogeny and the systematics of the Gastropoda is not yet clear.     

Geological history and phylogeny of Chelicerata

Chelicerata probably appeared during the Cambrian period. Their precise origins remain unclear, but may lie among the so-called great appendage arthropods. By the late Cambrian there is evidence for both Pycnogonida and Euchelicerata. Relationships between the principal euchelicerate lineages are unresolved, but Xiphosura, Eurypterida and Chasmataspidida (the last two extinct), are all known as body fossils from the Ordovician. The fourth group, Arachnida, was found monophyletic in most recent studies. Arachnids are known unequivocally from the Silurian (a putative Ordovician mite remains controversial), and the balance of evidence favours a common, terrestrial ancestor. Recent work recognises four principal arachnid clades: Stethostomata, Haplocnemata, Acaromorpha and Pantetrapulmonata, of which the pantetrapulmonates (spiders and their relatives) are probably the most robust grouping. Stethostomata includes Scorpiones (Silurian–Recent) and Opiliones (Devonian–Recent), while Haplocnemata includes Pseudoscorpiones (Devonian–Recent) and Solifugae (Carboniferous–Recent). Recent works increasingly favour diphyletic mite origins, whereby Acaromorpha comprises Actinotrichida (Devonian–Recent), Anactinotrichida (Cretaceous–Recent) and Ricinulei (Carboniferous–Recent). The positions of the Phalangiotarbida (Devonian–Permian) and Palpigradi (Neogene–Recent) are poorly resolved. Finally, Pantetrapulmonata includes the following groups (listed here in their most widely recovered phylogenetic sequence): Trigonotarbida (Silurian–Permian), Uraraneida (Devonian–Permian), Araneae (Carboniferous–Recent), Haptopoda (Carboniferous), Amblypygi (?Devonian–Recent), Thelyphonida (Carboniferous–Recent) and Schizomida (Paleogene–Recent).     

Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

The Paleozoic evolution of the gastropod larval shell: larval armor and tight coiling as a result of predation‐driven heterochronic character displacement

Early and middle Paleozoic gastropod protoconchs generally differ strongly from their corresponding adult morphologies, that is, most known protoconchs are smooth and openly coiled, whereas the majority of adult shells are ornamented and tightly coiled. In contrast, larval and adult shells of late Paleozoic gastropods with planktotrophic larval development (Caenogastropoda, Neritimorpha) commonly resemble each other in shape and principle ornamentation. This is surprising because habitat and mode of life of planktonic larvae and benthic adults differ strongly from each other. Generally, late Paleozoic to Recent protoconchs are tightly coiled. This modern type of larval shell resembles the adult shell morphology and was obviously predisplaced onto the larval stage during the middle Paleozoic. The oldest known planktonic‐armored (strongly ornamented) larval shells are known from the late Paleozoic. However, smooth larval shells are also common among the studied late Paleozoic gastropods. The appearance of larval armor at the beginning of the late Paleozoic could reflect an increase of predation pressure in the plankton. Although there are counter examples in which larval and adult shell morphology differ strongly from each other, there is statistical evidence for a heterochronic predisplacement of adult characters onto the larval stage. Larval and adult shells are built in the same way, by accretionary secretion at the mantle edge. It is likely that the same underlying gene expression is responsible for that. If so, similarities of larval and adult shell may be explained by gene sharing, whereas differences may be due to different (planktic vs. benthic life) epigenetic patterns.     

Paleozoic bryozoans of Mongolia

Stratigraphic assemblages characteristic for the Ordovician, Silurian, Devonian, Carboniferous, and Permian are presented based on the analysis of newly refined and supplemented data on the composition and distribution of Paleozoic bryozoans in Mongolia. Thirty-four auxiliary biostratigraphic units ranked as beds with bryozoans are established in order to subdivide the Paleozoic strata of Mongolia.     

Composition and dynamics of the great Phanerozoic Evolutionary Floras

Factor analysis of a data set representing the global distribution of vascular plant families through time shows the broad pattern of vegetation history can be explained in terms of five Evolutionary Floras. The Rhyniophytic (=Eotrachyophytic) Flora represents the very earliest (Silurian and earliest Devonian) vascular plants, notably the Rhyniophytopsida. The Eophytic Flora represents the early (Early–Middle Devonian) mainly homosporous land plants, notably the Zosterophyllopsida, Trimerophytopsida and early Lycopsida. The Palaeophytic Flora represents the Late Devonian and Carboniferous vegetation, which saw the introduction of heterospory among the spore producing plants and of early gymnosperms. The Mesophytic Flora first appeared in the Late Carboniferous and Permian macrofossil record, although there is palynological evidence of these plants having grown earlier in extra‐basinal habitats and was dominated by gymnosperms with more modern affinities. The Cenophytic Flora that first appeared during Cretaceous times was overwhelmingly dominated by angiosperms. The end‐Devonian, end‐Triassic and end‐Cretaceous mass‐extinction events recognized in the marine fossil record had little impact on the diversity dynamics of these Evolutionary Floras. Rather, the changes between floras mainly reflect key evolutionary innovations such as heterospory, ovules and angiospermy.     

Early evolutionary trends in ammonoid embryonic development

During the Devonian Nekton Revolution, ammonoids show a progressive coiling of their shell just like many other pelagic mollusk groups. These now extinct, externally shelled cephalopods derived from bactritoid cephalopods with a straight shell in the Early Devonian. During the Devonian, evolutionary trends toward tighter coiling and a size reduction occurred in ammonoid embryonic shells. In at least three lineages, descendants with a closed umbilicus evolved convergently from forms with an opening in the first whorl (umbilical window). Other lineages having representatives with open umbilici became extinct around important Devonian events whereas only those with more tightly coiled embryonic shells survived. This change was accompanied by an evolutionary trend in shape of the initial chamber, but no clear trend in its size. The fact that several ammonoid lineages independently reduced and closed the umbilical window more or less synchronously indicates that common driving factors were involved. A trend in size decrease of the embryos as well as the concurrent increase in adult size in some lineages likely reflects a fundamental change in reproductive strategies toward a higher fecundity early in the evolutionary history of ammonoids. This might have played an important role in their subsequent success as well as in their demise.     

ORIGIN, EVOLUTION AND CLASSIFICATION OF THE NEW SUPERORDER NEPIOMORPHIA (MOLLUSCA, BIVALVIA, LOWER PALAEOZOIC)

Abstract: A number of bivalve taxa defined in the past as ‘Cryptodonten’ by Neumayr, 1884 were grouped together in the high‐level taxon Palaeoconchae Neumayr, 1884. Cox (1969) noted that Palaeoconchae and Cryptodonta were synonymous and Newell (1969) used Cryptodonta as a subclass of bivalves. However, for the past 120 years, the Cryptodonta has been poorly conceptualized and the name was used for poorly understood genera or those lacking dentition. As used by Newell, Cryptodonta included taxa now placed in the subclasses Protobranchia Pelseneer and Autolamellibranchiata Grobben, and to the class Rostroconchia Pojeta, Runnegar, Morris and Newell. The bulk of Newell's use of Cryptodonta was made up of Silurian and Devonian taxa first described by Barrande (1881) from Bohemia; Newell placed these in the order Praecardioida Newell. In effect, Cryptodonta became a ‘wastebasket’ grouping for what, at the time, were poorly understood taxa. Many of the formerly poorly understood praecardioids are now better known and are herein placed in the new superorder Nepiomorphia. The Nepiomorphia contains two orders: (1) order Praecardioida that includes the families Slavidae K?í?, Cardiolidae Hoernes, Praecardiidae Hoernes and Buchiolidae Grimm; and (2) new order Antipleuroida that includes the families Stolidotidae fam. nov., Spanilidae fam. nov. and Antipleuridae Neumayr. The Nepiomorphia originated probably in the early Silurian as result of r‐selection progenesis. When the marine current system became re‐established after the late Ordovician glaciation and in the early Silurian, an at least temporary ventilation of the shallow waters by surface currents was renewed in higher latitudes of peri‐Gondwana and Siberia, producing acceptable sea‐bottom environments. Larvae were distributed by surface currents from the warm tropical regions of Laurentia and Baltica and were among the first benthic organisms to colonize the new environments. Temporary ventilation created frequent density‐independent catastrophic mortalities of early ontogenetic stages, with no competitors and with super‐abundant resources. During the Silurian and early Devonian, the Nepiomorphia underwent several diversifications in the recurring cephalopod limestone biofacies characteristic of peri‐Gondwana, and evolved infaunal, semi‐infaunal and epifaunal modes of life in several lineages. The Nepiomorphia most probably became extinct during the early Carboniferous and had no role in the future evolution of the Bivalvia.     

Studies of fossil brachiopods of Mongolia: Ordovician,Silurian, Devonian,Carboniferous, and Permian

Fossil brachiopods from the Ordovician, Silurian, Devonian, Carboniferous, and Permian deposits of Mongolia have been studied for the last forty-five years by the Joint Soviet-Mongolian (later RussianMongolian) Paleontological and Geological Expeditions. New data on the taxonomic composition, stratigraphic and geographic distribution of the brachiopod assemblages have been obtained. The brachiopod systematics has been further refined and detailed, and the stratigraphic and correlation scales and biogeographic reconstructions have been elaborated for the Paleozoic of Mongolia.     

DEVONIAN CONODONT APPARATUSES AND THEIR VICARIOUS SKELETAL ELEMENTS

Recognition of four types of skeletal apparatuses of Devonian compound and platform conodonts, which conform in their basic plan to those already established in the Ordovician, Silurian, and Carboniferous, emphasizes the persistence of but a few structural types in the Paleozoic. The vicarious nature primarily of compound but also of platform elements in different multielement associations demonstrates mosaic evolution in Devonian apparatuses and has far-reaching taxonomic implications. Homologies of skeletal elements, for which a system of symbols is introduced, are evident in different apparatus types and show that the relationships of elements have been obscured by the older form taxonomy. Multielement analysis revises the taxonomic scope of the common Devonian genera, Icriodus, Polygnathus , and Ozarkodina. As a consequence of apparatus reconstruction, four new genera are proposed: Parapolygnathus, Cryptotaxis, Delotaxis , and Pedavis.     

On the origin of bactritoids (Cephalopoda)

There is a high probability that bactritoids represent a paraphylum or polyphylum. The initial chambers or protoconchs of the Early-Middle DevonianBactrites Sandberger,Devonobactrites Shimansky, andLobobactrites Schindewolf are elongated spheres with a diameter of 0.3–1.0 mm. The initial chambers are larger in diameter than the slender, smooth shaft located adorally to the initial chamber. Similar apices occur in a number of Late Silurian sphaerorthoceridans with central siphuncles. Sphaerorthoceridans with a bactritoid-like apex and an eccentric siphuncle are known from the Early Devonian. The earliest questionableBactrites occurs in the Pragian (middle Early Devonian). By Emsian time bactritoids are common elements of cephalopod faunas.Bactrites-like orthocones of the Middle Ordovician and Late Silurian are homeomorphs with clearly different early growth stages. Thus, the time interval between the first appearance ofBactrites and the origin of ammonoids can be narrowed down to the Pragian to Early Emsian. The placement of the siphuncle in a ventral marginal position has been used as one of the critical morphologic features in defining the bactritoids. However, the displacement of the siphuncle from subcentral or eccentric positions toward the conch margin occurred at least three times during the Ordovician — Early Devonian evolution of the Orthocerida. Thus, there is a high probability that a marginal shift of the orthocerid siphuncle occurred in post-Emsian times, too.     

Evolutionary and biogeographical shifts in response to the Late Ordovician mass extinction

The Late Ordovician mass extinction was an interval of high extinction with inferred low ecological selectivity, resulting in little change in community structure after the event. In contrast, the mass extinction may have fundamentally changed evolutionary dynamics in the surviving groups. We investigated the phylogenetic relationships among strophomenoid brachiopods, a diverse brachiopod superfamily that was a primary component of Ordovician ecosystems. Four Ordovician families/subfamilies sampled in the analysis (Rafinesquinidae, Strophomeninae, Glyptomenidae and Furcitellinae) were reconstructed as monophyletic groups, and the base of the strophomenoid clade that dominated the Silurian recovery was reconstructed as diversifying alongside these families during the Middle Ordovician. We time‐calibrated the phylogeny and used geographical occurrences to investigate biogeographical changes in the strophomenoids through time with the R package BiogeoBEARS . Our results indicate that extinction was higher in taxa whose ranges were constrained to tropical or subtropical regions. Furthermore, our results suggest important shifts in the diversification patterns of these brachiopods after the mass extinction. While most of the strophomenoid families survived the Late Ordovician event, ecologically abundant taxonomic groups during the Ordovician were either driven to extinction, reduced in diversity, or slowly died off during the Silurian. The new abundant strophomenoid taxa derived from one clade (consisting of Silurian–Devonian groups such as Douvillinidae, Strophodontidae and Amphistrophiidae) that diversified during the post‐extinction radiation. Our results suggest the selective diversification during the Silurian radiation, rather than selective extinction in the Late Ordovician, had a greater impact on the evolutionary history of strophomenoid brachiopods.     

Palaeozoic Foraminifera: Systematics, palaeoecology and responses to global changes

Because the Foraminifera are very sensitive to various environmental parameters (e.g., water temperature, salinity, light, etc.), there are important proxies used for palaeoenvironmental and palaeogeographic reconstructions. The evolution of the structure, shape and size of the mineralized tests of Foraminifera can directly reflect the variation of these parameters through geological time. Furthermore, their biostratigraphic value has been widely demonstrated. In this context, the systematics, evolution and ecological behaviour of the first mineralized Palaeozoic Foraminifera are important to discuss in order to have a clearer picture of former shallow marine environments, and finally understand their distribution through space and time. The systematics of the fossil group of Foraminifera that first developed a mineralized test remains under discussion. These early foraminifers are considered as Textulariata (as generally admitted), recrystallized Fusulinata or an independent group, sometimes called Astrorhizata. In this paper, we argue to assign the early foraminifers to the Fusulinata, and to subdivide this class into six orders: Parathuramminida, Archaediscida and Earlandiida (forming together the subclass Afusulinana n. subcl.), and Tournayellida, Endothyrida and Fusulinida (subclass Fusulinana nom. translat.). These subdivisions are discussed and linked to the first occurrences of the later classes: Miliolata, Nodosariata and Textulariata. The environmental living conditions of the first fossilized foraminifers remain enigmatic during the Early Palaeozoic (Cambrian-Silurian). During the Late Silurian, the unilocular Parathuramminida started to colonize the inner parts of ramps and platforms. The first plurilocular microgranular foraminifers (Semitextulariidae, Nanicellidae, and Eonodosariidae) developed in back-reefal systems and in deeper-water environments (“griottes”-type nodular limestone) from the late Early Devonian to the early Late Devonian. The Moravamminida, another group of possible Protista, are typical markers of Devonian inner ramp systems. The Semitextulariidae, Nanicellidae, and Eonodosariidae did not survive the Frasnian/Famennian crisis. From the Tournaisian to the Serpukhovian (Mississippian subsystem or Early Carboniferous), numerous new genera and species of Archaediscida, Tournayellida and Endothyrida flourished but remained confined to inner ramp environments. In deeper water depositional systems (i.e. coral thrombolite microbialites and/or nodular limestones), a few opportunistic Foraminifera were living up to the disphotic zone. During the Pennsylvanian (Bashkirian to Gzhelian), the habitats extended to more confined, shallower areas of the inner ramp (with Staffelloidea). During the Late Carboniferous and Permian, the larger Fusulinida (Schwagerinoidea) reached the outer platform as they have been commonly reworked in calciturbidites. During the Late Permian, some taxa were even able to live in hypersaline environments such as sabkhas and hypersaline lagoons. Two major biotic crises occurred during the Permian (post-Middle and post-Late Permian crisis), but the number of survivors after the PTE (Permian/Triassic Extinction) is probably higher than previously admitted. From the Cambrian to the Serpukhovian, the Foraminifera were probably all infaunal or living at the sediment/seawater interface. The TROX and TROX-2 models are consequently applicable. Anoxia, often suggested as triggering environmental crises, was likely not systematically lethal for many infaunal foraminifers. The late Tournaisian-Changhsingian Tetrataxis genus was probably the first epiphyte foraminifer, because of its conical, limpet-like test. The Tetrataxidae (e.g., Tetrataxis, Pseudotaxis and Abadehella) constituted the unique trochospirally coiled plurilocular foraminiferal family of the Palaeozoic. The Bradyinoidea, Ozawainelloidea, Staffelloidea, and the Pseudoschwagerinidae (Schwagerinoidea) are other examples of Pennsylvanian-Permian epiphytes but cannot be considered as planktonic taxa. All the other Schwagerinoidea are related to high-energy environments and coarse-grained substrates. Their history, as well as that of the Neoschwagerinoidea, was likely subject to the vicissitudes of their endosymbiotic algae.     

Microbrachiopods and the end‐Ordovician event

Millimetre sized chitinophosphatic brachiopods ("microbrachiopods") largely, but by no means entirely, centred around the family Acrotretidae are commonly regarded as being in decline after the early Ordovician. Work on Irish Upper Ordovician limestones however shows that this is not the case, material recovered showing both numerical abundance and taxonomic diversity in beds of Ashgill age. Although forms are known from the Devonian, the published record of these neglected fossils from the Silurian is sparse, so that the effects, if any, of the end‐Ordovician event on this ecologically enigmatic group cannot as yet be determined.     

Phanerozoic changes in the high-rank suprageneric diversity structure of brachiopods: Linear and non-linear effects

Phanerozoic evolution of brachiopods produced many linear (established by a comparison of successive geologic time units) and non-linear (established by a comparison of non-successive geologic time units) effects, which can be examined quantitatively by using the similarity coefficients (Czekanowski's Quantified Coefficient and Gower Index) and correlation tools. The high-rank suprageneric diversity structure accounts for a number of superfamilies in each of 26 orders for every epoch of geological time. The intensity of turnovers in this structure was generally low during the entire Phanerozoic. It was slightly stronger during the Early Paleozoic, but close to zero during the Cenozoic, when the high-rank suprageneric diversity structure of brachiopods stabilized finally. Significant turnovers took place at the Middle Cambrian–Early Ordovician, the Late Ordovician–Early Silurian, the Late Silurian–Early Devonian, the Middle Devonian–Mississippian, and the Permian–Triassic transitions. Influences of mass extinctions, both major like those End Ordovician or Permian/Triassic and minor like Early Jurassic or Jurassic/Cretaceous, on the high-rank suprageneric diversity structure of brachiopods is registered. The strongest was the consequences of the Permian/Triassic catastrophe, which perhaps even reset the brachiopod evolution. No evident direct relationships are established between intensity of turnovers and eustatic fluctuations. However, the changes in the diversity structure recorded with the Gower Index provide evidence that eustatic lowstands were more favorable for intensification in these changes.     

First Mesozoic Scutigeromorph Centipede, from the Lower Cretaceous of Brazil

The first Mesozoic scutigeromorph centipede (Myriapoda: Chilopoda), Fulmenocursor tenax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. Previously described fossil Scutigeromorpha are known from Dominican and Baltic amber, the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois, the Silurian and Devonian of Britain, and the Devonian of New York State.     

Latest Helcionelloid Molluscs from the Lower Ordovician of Kazakhstan

The helcionelloid mollusc Chuiliella elenae gen. et sp. nov. is described from the Lower Ordovician of Kazakhstan. It represents the geologically youngest record of a group of mainly bilaterally symmetrical ancient molluscs which originated in the earliest Cambrian, flourished during the early–mid Cambrian and was thought to have become extinct during the late Cambrian. Chuiliella is a typical helcionelloid in terms of shell shape, although the comarginal ornamentation characteristic of many helcionelloids is lacking. Interpretation of the raised margin of the aperture adjacent to the earlier coiled whorl as exhalant channels favours reconstruction of helcionelloids as endogastrically coiled, i.e., with the apex posterior.     

Did the Late Ordovician mass extinction event trigger the earliest evolution of ‘strophodontoid’ brachiopods?

‘Strophodontoid’ brachiopods represented the majority of strophomenide brachiopods in the Silurian and Devonian periods. They are characterized by denticles developed along the hinge line. The evolution of denticles correlated with the disappearance of dental plates and teeth and were already present when the clade originated in the Late Ordovician. Specimens of Eostropheodonta parvicostellata from the Kuanyinchiao Bed (early–middle Hirnantian, uppermost Ordovician) in the Hetaoba Section, Meitan, Guizhou Province, South China, display clear fossil population variation, during a process of loss of dental plates and the development of denticles. Three phenotypes of E. parvicostellata are recognized in a single fossil bed, likely heralding a speciation process. Non-metric multidimensional scaling (NMDS) based on five key characters of genera of the Family Leptostrophiidae shows a much wider morphospace for Silurian genera than for those in the Devonian. Phylogenetic analysis of the Family Leptostrophiidae supports the NMDS analysis and mostly tracks their geological history. The fossil population differentiation in E. parvicostellata discovered between the two phases of the Late Ordovician mass extinction event (LOME) linked to a major glaciation, suggests a Hirnantian origination of the ‘strophodontoid’ morphology, and links microevolutionary change to a macroevolutionary event.     

The development of early terrestrial ecosystems

Summary

In this review of terrestrialization by plants and animals in the early Phanerozoic, the classical idea of a major mid-Palaeozoic event is discarded in favour of gradual colonization over a long time period. Four phases of colonization of the land by plants are recognized but their limits are often difficult to define. The first, of microbial mats comprising prokaryotes and later photosynthesizing protists (algae) but with no direct fossil evidence, extends from the Precambrian and may persist in environments unsuitable for colonization by higher plants and animals today. The second, based on microfossils (spores and cuticles) possibly from plants of bryophyte aspect (if not affinity) started in the Ordovician (c. 460 Ma ago) and ended in the Lower Devonian, but was overlapped by the third phase beginning early in the Silurian (c. 430 Ma). This consisted of small plants of axial organization with terminal sporangia probably allied to the tracheophytes. The advent of taller vascular plants of varied organization around the Silurian — Devonian boundary (c. 420–400 Ma) signalled the final pioneering phase — that of major adaptative radiations on land, culminating in the appearance of extant groups, in changes in reproductive strategy and in the development of complex vegetation structure. The animal record is sparser than that of the plants, but suggests an early land fauna in the mid-Palaeozoic which differed from later terrestrial assemblages in lacking herbivores, with the first direct fossil evidence for land animals in the late Silurian.