Rise and fall of the phacopids: the morphological history of a successful trilobite family

Phacopidae were a successful family of the Silurian–Devonian period. Although their diversity trends are well identified, their shape evolution is unknown; their morphology often considered to be conservative. We have quantified these morphologies using geometric morphometrics (landmarks) and investigated their evolution using morphological disparity indices. Results identified morphological variations between the genera, and through time. Phacopids differ from each other by the position of the facial suture linked to the size of the visual complex, the shape of the genal angle and the elongation of both cephalon and pygidium. The morphological disparity of cephala was high from the Silurian, contrary to that of pygidia. Subsequently, the morphological disparity increased in the Early Devonian with the development of narrow cephala and triangular pygidia. Morphological disparity was greater in the Emsian for both cephala and pygidia, more than 50 myr after the origination of phacopids. It constituted a perfect example illustrating that a peak of biodiversity does not necessarily happen in the early history of a clade. Subsequently, a strong decrease of morphological disparity occurred in the Middle Devonian, in conjunction with sea-level changes and anoxic events. Taxonomic richness and morphological disparity declined strongly in the Givetian, in a non-random extinction affecting particularly blind genera. The morphological disparity remained low in the Frasnian despite progressive eye reduction influenced by environmental changes. An extensive recovery occurred in the Famennian with an important increase of both taxonomic and morphological diversity. The Hangenberg event caused the final extinction of phacopids.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

澳大利亚泥盆纪皱纹珊瑚生物地层及生物地理研究展望

通过对澳大利亚泥盆纪皱纹珊瑚属级分布的分析,认为泥盆纪珊瑚在该区经历了两大繁盛期,尤以Ｌｏｃｈｋｏｖｉａｎ－Ｅｍｓｉａｎ最盛,有大量新属产生,内板珊瑚和菲力甫珊瑚科分子要Ｐｒａｇｉａｎ期开始出现,绳珊瑚科的分子也在Ｅｍｓｉａｎ大量出现,这些分子继而通过洋流逐渐扩散到华南及老世界域其他省区。     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Evolution of Devonian biodiversity of radiolarians with two porous spheres and one main spine

Features and major distinctions in the appearance and distribution of radiolarians with two porous spheres and one main spine in the Devonian basins are considered. Four solitary population waves and expansion scenario of radiolarians of this morphotype have been recognized in the Devonian. A new species, Radiobisphaera rozanovi sp. nov., from the Upper Emsian, Upper Eifelian, and Middle Frasnian of the southern Ural Mountains, from the Upper Eifelian of the Prague Basin (Barrandian), from the Middle-Upper Frasnian of the Rudnyi Altai, and from the Lower Famennian of the Timan-Pechora Basin and the Polar Ural Mountains is described.     

Famennian conodonts from the Hongguleleng Formation at the Bulongguoer stratotype section,western Junggar,Northwest China

The Hongguleleng Formation, the highest and most important Devonian marine carbonate horizon in western Junggar, contains an endemic shallow-water Icriodus–Polygnathus conodont fauna with rare palmatolepids and other genera. The conodont faunas from the Bulongguoer and the Wulankeshun sections are similar, with high abundance (about 40%) of endemic taxa (13 taxa), indicating isolation of the Junggar Basin during early Famennian. Non-endemic species in the faunas suggest that the Lower Member of the Hongguleleng Formation is assignable to the Pa. rhomboidea Zone to the Pa. marginifera marginifera Zone of early Famennian, not including the Frasnian–Famennian boundary. The Upper Member may be of late Famennian–early Tournaisian in age on the basis of our preliminary faunal analysis.     

Diversity dynamics of Zosterophyllopsida

The Zosterophyllopsida were major contributors to the diversification of early land plants. We present the first detailed analysis of the diversity dynamics of these plants from an updated database of all currently recognized zosterophyllopsid species. A set of quantitative methods classically used in palaeodiversity studies was applied to two data sets. The first one, ‘Zosterophyllopsida sensu stricto’, corresponds to the clade identified by Hao & Xue (The Early Devonian Posongchong Flora of Yunnan. (2013), Science Press). In the second, called ‘Zosterophyllopsida sensu lato’, barinophytalean‐type plants and taxa for which zosterophyllopsid affinities are suspected are added. The number of localities is used to explore sampling bias. Results show that sampling effect is minimal for the Early Devonian. For this time interval, both data sets record consistent patterns of changes suggesting that, whatever their affinities, all taxa included in the Zosterophyllopsida sensu lato show similar evolutionary trends. The diversity dynamics of zosterophyllopsids are characterized by a radiation during the Lochkovian, maximal values in the Pragian and a decline starting in the Emsian. The proportion of zosterophyllalean taxa with terminal sporangia is high until the Late Lochkovian when gosslingialean taxa without terminal sporangia evolved. During the Middle and Late Devonian, when diversity patterns are strongly affected by sampling, zosterophyllopsid diversity is low and characterized by a high proportion of barinophytacean and gosslingialean taxa, the latter becoming extinct in the Early Frasnian.     

Late Devonian global ostracod palaeobiogeography

A global Late Devonian ostracod database is constructed, incorporating new materials from South China and Northwest China. Four palaeobiogeographical units (Cathaysia, North America, Europe and peri‐Gondwana) are recognized during the Frasnian and five palaeobiogeographical units (Cathaysia, North America, Europe, Siberia and Australia) in the Famennian. Three controlling factors (climatic zonation, geographical isolation and global sea‐level changes) are identified to have played roles in shaping the palaeogeographical regionalization of ostracod faunas in the Late Devonian. The ostracod palaeobiogeography in the Frasnian was mainly influenced by climatic zonation, while rapid changes in tectonic configuration in the Famennian drastically altered the global palaeobiogeography of ostracods. The palaeobiogeographical regionalization of ostracod faunas suggests that Laurussia and Gondwana continued to draw near during the Late Devonian, with the first collision occurring in Southern Central Europe in the Famennian. The South China plate drifted northward to the Kazakhstan plate away from the Australian plate, which gradually became isolated during the Famennian Stage.     

Paléobiogéographie des faunes d'Ostracodes au Dévonien supérieur

About one hundred papers and studies have been utilized for a synthesis of ostracode palaeobiogeography during the Frasnian and Famennian. The pelagic entomozoid species have worldwide distributions: they are not used for distinguishing the palaeobiogeographic areas. Five provinces probably existed during the Frasnian and five during the Famennian. The palaeobiogeographic area of the Palaeotethys is constant during this time. There are important faunistic relations between the Western Canadian Platform and the Russian Platform, that is to say, along the Northern margin of the North-Atlantic Continent, during Late Devonian time. The province of Southeastern Asia remains isolated. The number and limits of the palaeobiogeographic areas have been controlled by climate and some geographical barriers.     

Extinctions and migrations of Devonian rugose corals in the Eastern Americas Realm

Oliver, W. A., Jr. 1990 04 15: Extinctions and migrations of Devonian rugose corals in The Eastern Americas Realm. Lethaia , Vol. 23 , pp. 167–178. Oslo. ISSN 0024–1164.
Detailed plotting of the stratigraphic ranges of Devonian rugose coral genera within the Eastern Americas Realm reveals new information about the extinctions and migrations of this largely endemic fauna. There were significant faunal turnovers in the Lochkovian, middle Eifelian and late Givetian. as well as in the often discussed late Frasnian. The late Givetian turnover was nearly as great as the Frasnian one. Inward migration was principally from western North America, and the greatest influxes were during the carly Givetian and Frasnian. It seems likely that there were several separate incursions and that some genera were introduced to the east two, or even three times.     

The late Devonian trilobite crises

Mid‐Devonian to end‐Late Devonian trilobites of different taxonomic categories are updated as to their actual stratigraphical range with respect to the internationally defined stage boundaries. The main palaeogeographical and ecological occurrences are summarized. Numerical analyses emphasize the clear relationship between fluctuations in diversity and global eustatic events. Already declining in diversity from the early mid‐Devonian, shallow‐water communities became most restricted during the mid‐Givetian Taghanic transgression. After a phase of adaptive radiation, off‐shore trilobite communities were severely affected during the mid‐ and end‐Late Devonian crises. From an initial 5 orders 3 were lost at the end‐Frasnian Kellwasser crisis while only 1 from the remaining 2 orders survived the Devonian‐Carboniferous boundary Hangenberg event. In both cases extinction was preceded by a unidirectional evolutionary trend in eye reduction accompanied by impoverishment of lower rank taxa. This phenomenon is obviously a result of selective adaptation under constant long‐lasting environmental influences. Specialization to obligate epi‐ or even endo‐benthic life habit, however, led fatally to extinction when stable conditions became substantially perturbed. Sudden sea‐level changes with subsequent break in the REDOX‐equilibrium took place at the Kellwasser and Hangenberg events, which were most probably responsible for trilobite mass extinctions.     

Significance of Devonian-Carboniferous radiolarians from accretionary terranes of the New England orogen, eastern Australia

Radiolarians provide age constraints for many previously undated terranes in the New England Orogen (NEO), a tectonic collage developed along the eastern margin of Australia.Djungati terrane, the age range of which was previously unknown, contains two distinctive siliceous sedimentary lithofacies. The oldest is a thick sequence of red, ribbon-bedded cherts which probably accumulated in a deep ocean-floor setting far from land. Middle Silurian through Late Devonian radiolarians have been recovered from these cherts. Green tuffaceous cherts which contain a latest Devonian (Famennian) radiolarian fauna depositionally overlie the lower red ribbon-bedded chert sequence. These cherts are intercalated with volcaniclastic sediments and the fauna which they contain can be used to constrain the timing of accretion of older rocks into a subduction complex.Anaiwan terrane, which was also previously undated, contains thin ribbon-bedded cherts which are depositionally overlain by tuffaceous chert, siliceous siltstones and volcaniclastic sediments. Latest Devonian (?late Famennian) and Early Carboniferous radiolarians have been recovered from these cherts and tuffaceous siltstones.Radiolarians also occur in fine-grained siliceous sediments of the Yarrimie Formation, part of the Gamilaroi terrane. These radiolarians are of Late Devonian (Frasnian) affinity and their presence indicates that blocks of limestone, which contain Givetian conodonts and corals and were previously thought to indicate the age of the Yarrimie Formation, are allochthonous.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.     

The oldest Gondwanan cephalopod mandibles (Hangenberg Black Shale,Late Devonian) and the mid‐Palaeozoic rise of jaws

It is widely accepted that the effects of global sea‐level changes at the transition from the Devonian to the Carboniferous are recorded in deposits on the shelf of northern Gondwana. These latest Devonian strata had been thought to be poor in fossils due to the Hangenberg mass extinction. In the Ma'der (eastern Anti‐Atlas), however, the Hangenberg Black Shale claystones (latest Famennian) are rich in exceptionally preserved fossils displaying the remains of non‐mineralized structures. The diversity in animal species of these strata is, however, low. Remarkably, the organic‐rich claystones have yielded abundant remains of Ammonoidea preserved with their jaws, both in situ and isolated. This is important because previously, the jaws of only one of the main Devonian ammonoid clades had been found (Frasnian Gephuroceratina). Here, we describe four types of jaws of which two could be assigned confidently to the Order Clymeniida and to the Suborder Tornoceratina. These findings imply that chitinous normal‐type jaws were likely to have already been present at the origin of the whole clade Ammonoidea, i.e. in the early Emsian (or earlier). Vertebrate jaws evolved prior to the Early Devonian origin of ammonoids. The temporal succession of evolutionary events suggests that it could have been the indirect positive selection pressure towards strong (and thus preservable) jaws since defensive structures of potential prey animals would otherwise have made them inaccessible to jawless predators in the course of the mid‐Palaeozoic marine revolution. In this respect, our findings reflect the macroecological changes that occurred in the Devonian. [Correction added on 28 July 2016 after first online publication: In the Abstract, the sentence “Vertebrate jaws probably … in the Early Devonian” was amended]     

Anatomical reinvestigation of Archaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Abstract Archaeopteris macilenta is one of the most widespread plants in the Late Devonian. Based on fossils from the Frasnian Huangjiadeng Formation, Yichang District of Hubei Province, for the first time we study in detail the anatomy of this progymnosperm plant in South China. Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue. Penultimate axes are eustelic, bearing eight sympodia and a thin band of secondary xylem. Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement. Archaeopteris macilenta and A. halliana (A. roemeriana) are dominant in the Frasnian and Famennian, respectively. Comparisons with these two species from other tectonic plates indicate consistent stelar architectures. Global spread, continuous occurrence, and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian–Famennian extinction event. In this time, endemic genera and cosmopolitan taxa, including Archaeopteris, suggest the palaeogeographic isolation of South China and certain associations with other plates.     

Contrasting environmental and regional effects on global pteridophyte and seed plant diversity

Pteridophytes (ferns and fern‐allies) represent the second‐largest group of vascular plants, but their global biogeography remains poorly studied. Given their functional biology, pteridophytes are expected to show a more pronounced relation to water availability and a higher dispersal ability compared to seed plants. We test these assertions and document the global pattern of pteridophyte richness across 195 mainland and 106 island regions. Using non‐spatial and spatial simple and multiple regression models, we analyze geographic trends in pteridophyte and seed plant richness as well as pteridophyte proportions in relation to environmental and regional variables. We find that pteridophyte and seed plant richness are geographically strongly correlated (all floras: r=0.68, mainland: r=0.82, island floras: r=0.77), but that the proportions of pteridophytes in vascular plant floras vary considerably (0–70%). Islands (mean=15.3%) have significantly higher proportions of pteridophytes than mainland regions (mean=3.6%). While the relative proportions of pteridophytes on islands show a positive relationship with geographic isolation, proportions in mainland floras increase most strongly along gradients of water availability. Pteridophyte richness peaks in humid tropical mountainous regions and is lowest in deserts, arctic regions, and on remote oceanic islands. Regions with Mediterranean climate, outstanding extra‐tropical centres of seed plant richness, are comparatively poor in pteridophytes. Overall, water‐energy variables and topographical complexity are core predictors of both mainland pteridophyte and seed plant richness. Significant residual richness across biogeographic regions points to an important role of idiosyncratic regional effects. Although the same variables emerge as core predictors of pteridophyte and seed plant richness, water availability is clearly a much stronger constraint of pteridophyte richness. We discuss the different limitations of gametophytes and sporophytes that might have limited the ability of pteridophytes to extensively diversify under harsh environmental conditions. Our results point to an important role of taxon‐specific functional traits in defining global richness gradients.     

Relative importance of water, energy, and heterogeneity in determining regional pteridophyte and seed plant richness in China

Environmental variables, such as ambient energy, water availability, and environmental heterogeneity have been frequently proposed to account for species diversity gradients. How taxon-specific functional traits define large-scale richness gradients is a fundamental issue in understanding spatial patterns of species diversity, but has not been well documented. Using a large dataset on the regional flora from China, we examine the contrast spatial patterns and environmental determinants between pteridophytes and seed plants which differ in dispersal capacity and environmental requirements. Pteridophyte richness shows more pronounced spatial variation and stronger environmental associations than seed plant richness. Water availability generally accounts for more spatial variance in species richness of pteridophytes and seed plants than energy and heterogeneity do, especially for pteridophytes which have high dependence on moist and shady environments. Thus, pteridophyte richness is disproportionally affected by water-related variables; this in turn results in a higher proportion of pteridophytes in regional vascular plant floras (pteridophyte proportion) in wet regions. Most of the variance in seed plant richness, pteridophyte richness, and pteridophyte proportion explained by energy is included in variation that water and heterogeneity account for, indicating the redundancy of energy in the study extent. However, heterogeneity is more important for determining seed plant distributions. Pteridophyte and seed plant richness is strongly correlated, even after the environmental effects have been removed, implying functional linkages between them. Our study highlights the importance of incorporating biological traits of different taxonomic groups into the studies of macroecology and global change biology.     

华南泥盆系法门阶浅海相四射珊瑚的组合序列

晚泥盆世弗拉期末F-F的生物灭绝事件,导致珊瑚以及许多浅海底栖生物灭绝,几乎整个法门期珊瑚群都处于残存阶段,华南地区目前只在湖南的个别地方发现少量的Smithiphyllum。直到法门期末珊瑚和其它一些造礁底栖生物开始复苏,出现了不少与典型泥盆纪珊瑚存在很大差别的新分子。华南泥盆纪最晚期(Strunian)的珊瑚可划分成上、下两个组合:上部Cystophrentis组合;下部Eocaninophyllum组合。泥盆—石炭纪之交的另一次生物灭绝事件,使新生的泥盆纪最晚期的珊瑚又遭灭绝。