Patterns of evolution and extinction in proetid trilobites during the late Devonian mass extinction event,Canning Basin,Western Australia

Abstract: In the early Late Devonian, terminal Frasnian proetid trilobites have previously only been known from Europe and North Africa. For the first time, a rich fauna of late Frasnian proetids is described from the Virgin Hills Formation, Canning Basin, Western Australia. Seventeen species in six genera are described, of which three are new: Rudybole gen. nov., Palpebralina gen. nov. and Canningbole gen. nov. A new subgenus, Chlupaciparia (Australoparia) subgen. nov. is also described. Fourteen of the species are new: Palpebralia initialis sp. nov., P. pustulata sp. nov., ?P. sp. nov. A, Rudybole depressa sp. nov., Palpebralina pseudopalpebralis sp. nov. (comprising the subspecies P. pseudopalpebralis pseudopalpebralis subsp. nov. and P. pseudopalpebralis ultima subsp. nov.), P. minor sp. nov., P. ocellifer sp. nov., Canningbole latimargo sp. nov., C. henwoodorum sp. nov., C. macromma sp. nov., Pteroparia extrema sp. nov., Chlupaciparia (Chlupaciparia) planiops sp. nov., Chlupaciparia (Australoparia) australis sp. nov. and C. (Australoparia) lata sp. nov. The subspecies Rudybole adorfensis angusta subsp. nov. is also described. The proetids range through conodont Zones 11–13b and terminate at the Upper Kellwasser Event, which marks the terminal Frasnian mass extinction event. Three of the six proetid lineages, Palpebralia, Canningbole and Pteroparia, show evolutionary trends of eye reduction. Two of the remaining lineages, Rudybole and Palpebralina, consist exclusively of blind taxa. The last, Chlupaciparia, also comprises forms with reduced eyes. The proetids show a stepped pattern of extinction during the late Frasnian, which correlate with two Kellwasser biocrises documented in European/North African Frasnian sections. The highest diversity preceded the Lower Kellwasser event that occurred at the end of conodont Zone 12 and saw the extinction of all species present in that zone. However, only one genus, Pteroparia, locally became extinct. A major higher‐level taxonomic mass extinction at the top of Zone 13b initiated the Upper Kellwasser extinction event. This included extinction at the generic level, with all five remaining genera becoming extinct, and at the family level, with the loss of the Tropidocoryphidae.     

PATTERNS OF EVOLUTION AND EXTINCTION IN THE LAST HARPETID TRILOBITES DURING THE LATE DEVONIAN (FRASNIAN)

Abstract: Late Devonian (Frasnian) harpetid trilobites have hitherto only been described from the western side of the Protethys Ocean, in what is now Europe and North Africa, as well as from Gondwana‐derived northwestern Kazakhstan (Mugodjar). However, late Frasnian strata in the Canning Basin, Western Australia, that were deposited on the eastern side of this ocean, contain a rich harpetid fauna. Described herein are two new harpetids: Eskoharpes gen. nov. and Globoharpes gen. nov., within which are placed six species: E. palanasus sp. nov., E. wandjina sp. nov., E. boltoni sp. nov., E. guthae sp. nov., G. teicherti sp. nov. and G. friendi sp. nov. The ontogenetic development of E. palanasus, E. wandjina and G. teicherti are described, including the first unequivocal harpetid protaspis. Globoharpes exhibits evidence of sexual dimorphism in the development of a pronounced preglabellar boss in some specimens. This structure is thought to have functioned as a brood pouch. Such structures have previously only been described in Cambrian and Ordovician trilobites, and never before in harpetids. It is suggested that the characteristic harpetid fringe functioned as a secondary respiratory structure. The Eskoharpes lineage shows evolutionary trends that mirror changes seen in ontogenetic development of the youngest species, suggesting the operation of peramorphic processes. This is the first record of heterochrony in harpetids and the first documented example of peramorphosis in Devonian trilobites. These harpetids demonstrate a stepped pattern of extinction during the late Frasnian, probably related to the effects of the two Kellwasser biocrises that have been well documented in European Frasnian sections. Highly vaulted species of Eskoharpes and the strongly vaulted Globoharpes became extinct at the Lower Kellwasser Event. The flatter species of Eskoharpes became extinct at the base of the Upper Kellwasser Event shortly prior to the Frasnian/Famennian boundary. The extinction of these harpetids, along with contemporaneous forms from Europe, which are also discussed herein, marks the end of the trilobite order Harpetida worldwide.     

Sphaeroidal enrolment in middle Cambrian solenopleuropsine trilobites

Esteve, J., Zamora, S., Gozalo, R. & Liñán, E. 2010: Sphaeroidal enrolment in middle Cambrian solenopleuropsine trilobites. Lethaia, 10.1111/j.1502‐3931.2009.00205.x Fifty specimens belonging to species of Solenopleuropsis and Pardailhania from Spain and France demonstrate sphaeroidal enrolment in Cambrian trilobites for the first time. These solenopleuropsines show novel coaptative structures in different regions of the exoskeleton: in the cephalon there are vincular furrows and notches; in the thorax an articulating facet is developed at the pleural margins, with a ball and socket connection on the adaxial most portion, and an articulating half‐ring axially; the pygidium possesses an articulating facet. The interaction of these coaptative structures resulted in a sphaeroidal enrolment that was a progressive act from the first articulation between the occipital ring and the first segment to the pygidial articulating facet. A similar type of sphaeroidal enrolment is observed in the Devonian trilobite Phacops. Both Cambrian and Devonian trilobites developed a vincular furrow in the ventral surface of the cephalon to close their bodies tightly. In both cases, this is probably a convergent adaptation to protect against predators and obrution. Indeed, the enrolled trilobites are very common in obrution deposits restricted to shallow and soft muddy substrates. □Coaptative structures, convergence, Murero Formation, Pardailhania, Solenopleuropsinae, Solenopleuropsis.     

Sedimentary and faunal changes across the frasnian/famennian boundary in the canning basin of Western Australia

The Canning Basin of northwestern Australia is a key area for understanding global changes at the “Kellwasser Events” and the Frasnian‐Famennian boundary. Frasnian stromatoporoid‐coral‐cyanobacterial reef platforms stretched out for enormous distances along the palaeoshelf but in the early Famennian they were completely replaced by cyanobacterial reef platforms. An iridium anomaly in the sequence was formerly believed to be at or close to the boundary and was interpreted as possible evidence for an asteroid impact. Recent field work and detailed biostratigraphy in the area east and southeast of Fitzroy Crossing has given dating relevant to the timing and extent of sea level changes, hypoxic incursions and reef backstepping. Goniatites and conodonts provide correlations with the international biostratigraphy.

In the Horse Spring area the stage boundary falls within the Virgin Hills Formation which normally has a rich pelagic goniatite, nautiloid and conodont fauna. In the latest Frasnian (Zone 13 of Klapper 1989; regional Sphaeromanticoceras lindneri Zone) large allochthonous reef blocks moved downslope into the open marine basin. A diverse gastropod fauna is associated with the last atrypid brachiopods. The faunal record at the immediate boundary is obscured by dolomitisation but manticoceratid goniatites range into this level. There is no evidence for the organic‐rich dark Kellwasser limestone facies.

In the McWhae Ridge area two Frasnian goniatite horizons with Beloceras trilobites and the giant Mamicoceras guppyi and Sphaeromanticoceras lindneri transgress over the reef slope. Stromatolitic cyanobacterial beds mark condensations. Again there is no trace of the oxygen‐depleted Kellwasser facies. However, as at Horse Spring, manticoceratids persisted into dolomitic marker beds that have no other preserved macrofauna. The iridium anomaly associated with Frutexites postdates the Frasnian‐Famennian boundary and was formed by cyanobacterial concentration.     

Spores and the middle-upper Devonian boundary

The stratigraphic levels most favoured for the Middle-Upper Devonian boundary fall approximately within the range of the ammonoid Pharciceras lunulicosta Zone, i.e. from the Middle varcus Subzone to the base of the Lower asymmetricus Zone of the conodont scale. Spore data that are potentially useful for recognition of the boundary within this range have been correlated with conodont zones in marine facies in the Boulonnais region of France. A vast amount of information on spores from Middle-Upper Devonian boundary strata has accumulated in the European U.S.S.R., where the boundary is taken at a somewhat lower level. Late Givetian and early Frasnian continental strata of Melville Island in the Canadian Arctic contain species present in the Boulonnais or the European U.S.S.R., as well as species common to both regions. Diatomozonotriletes spp., Rhabdosporites langii, Samarisporites triangulatus, Contagisporites optivus, Archaeoperisaccus timanicus, Chelinospora concinna and Ancyrospora langii, among others, may be useful for correlating the boundary as eventually defined. The stratigraphic ranges of most of these taxa show only limited agreement interregionally at present, probably owing at least in part to problems of spore nomenclature and taxonomy, and an insufficiency of spore reference sequences keyed to faunal zones. Nevertheless, individual species of spores, and especially assemblages of species, have much potential for delimiting and correlating the Middle-Upper Devonian boundary in both marine and continental facies.     

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.     

Frasnian conodonts from the eastern Russian Platform

The Frasnian conodonts of the eastern Russian Platform from the Timan in the north to the Orenburg Region in the south are characterized in detail both paleontologically and stratigraphically. This paper is a continuation of a previously published monograph (Ovnatanova and Kononova, 2001) on the Frasnian conodonts of the shallow-water strata of the central Russian Platform, where polygnathids prevail. The conodont assemblages from the shallow-water and basinal sections of the Volga-Ural province and Southern Timan are analyzed. Problems and difficulties associated with the correlation of the shallow-water and deep-water Frasnian sections of the eastern Russian Platform and their correlation with the existing zonal conodont scales are discussed. The correlation between the deep-water Mendym deposits and the shallow-water Rechitza and Voronezh strata of the Volga-Ural province with the Vetlasyan and Sirachoi deposits of the Southern Timan is substantiated. The stratotype of the Semiluki Regional Stage of the Central Devonian Field contains equivalents of the lowermost beds of the Domanik Formation (unit 1), based entirely on polygnathids. Phylogenetic reconstructions for Palmatolepis and Polygnathus are suggested based on the ontogenetic series for some species of these two genera and the presence of transitional forms between some of the species. Based on these phylogenetic reconstructions, conodont zonal scales for the shallow-water and deep-water sections of the Frasnian Stage of the Russian Platform are proposed and their correlation with the existing conodont zonal scales is also adduced. Some aspects of biofacies control are considered based on the distribution of conodonts in the sections studied. In Systematic Paleontology, 91 conodont species of the genera Ancyrodella, Mesotaxis, Palmatolepis, and Polygnathus are described, including the new species Palmatolepis menneri, P. kaledai, P. acutangularis, and Polygnathus reitlingerae. DEDICATION To the blessed memory of the outstanding geologist Vladimir Vladimirovich Menner     

Biostratinomy and functional morphology of enrollment in two Middle Devonian trilobites

Although it is common knowledge that many trilobites enrolled, behavioral and functional aspects of enrollment are not at all well understood. Taphonomic details portrayed by enrolled trilobites in the Middle Devonian Hamilton Group (New York State) indicate that enrollment was a complex and morphologically constrained behavior. The trilobites Phacops rana (Green) and Greenops boothi (Green) are frequently enrolled in Hamilton strata; biostratinomic data indicate two very different enrollment postures. Interlocking morphologies (coaptative devices) and apodeme structure and disposition indicate that these postures reflect specific behaviors which involved interaction between tergal structures, inferred musculature, and the substratum. Phacops enrolled by burrowing forward and down into the sediment; dorsal muscles, attached to prominent articulating half-rings, imbricated the thorax such that each lappet overlapped the next posterior segment and locked into a posterior pleural facet. The pygidium was brought into place as the posterior segments of the thorax were placed into vincular notches along the lateral margin of the ventral cephalon. The pygidium locked with the cephalic vincular furrow to complete ‘perfect sphaeroidal’ closure. Greenops enrolled with the cephalon in an upright position at the sediment surface; a submarginal furrow on the ventral surface of the pygidium received the anterior rim of the cephalon. Relatively narrow articulating half-rings limited pleural rotation. Segments were loosely locked into narrow facets at the anterior margin of the next posterior lappet. In spite of rudimentary lappet and half-ring structures, Greenops displays an elaborate system of thoracopygidial muscles which involved dorsoventral and longitudinal attachments along the thorax and into the pygidium. Phacops, in contrast, displays very poorly developed apodemes which occur in the thorax only. Longitudinal muscle strength was likely less important during Phacops enrollment than is evident for the Greenops enrollment procedure. Conversely, Phacops clearly relied to a great degree upon competent closure devices which are poorly developed in Greenops. Biostratinomic data reveal different enrollment behaviors which reflect the function of different enrollment-related morphologies present in each species.     

新疆准噶尔盆地西北缘洪古勒楞组时代的新认识

标准地点的洪古勒楞组长期以来一直被认为是准噶尔盆地西北缘,乃至整个新疆北部以海相为主的层位上最高的晚泥盆世晚期的沉积。然而牙形类和微体脊椎动物的研究证实,该组跨弗拉阶－法门阶（Ｆ－Ｆ）界线,它至少包括了一小部分弗拉晚期和主体部分为法门早期的沉积。两个牙形类带ＬａｔｅｒｈｅｎａｎａＺｏｎｅ和ＭｉｄｄｌｅｃｒｅｐｉｄａＺｏｎｅ被识别出,它们在Ｆ－Ｆ界线附近,并分别在大绝灭事件（在ｌｉｎｇｕｉｆｏｒｍｉｓＺｏｎｅ内）之前后。另一个层位更高的牙形类带ＬａｔｅｃｒｅｐｉｄａＺｏｎｅ也可能存在。同时还证实,长期以来一直被认为“正常层序”的标准地点的洪古勒楞组的层序是倒转的。由此,含植物大化石Ｌｅｐｔｏｐｈｌｏｅｕｍｒｈｏｂｉｃｕｍ等的陆相层在下（此层可能归于该组下伏的朱鲁木特组）,而产牙形类和其它丰富的无脊椎动物的海相层在上。按牙形类年代带的年代地层时间表（Ｆｏｒｄｈａｍ,１９９２）,该组距今约３６８－３６４百万年。至于它和邻近地区和布克河组的关系,由于后者发现了更高层位的牙形类,两者还不能完全对比。     

Progressive drowning of carbonate platform in the Moravo-Silesian Basin (Czech Republic) before the Frasnian/Famennian event: facies,compositional variations and gamma-ray spectrometry

The Moravo-Silesian Basin (MSB; eastern Czech Republic and southern Poland) hosted an extensive shallow-water carbonate platform in the Middle Devonian to Frasnian interval. The platform drowned in a stepwise fashion from the Palmatolepis hassi to the Pa. linguiformis zone. Three types of drowning successions were revealed from conodont biostratigraphy, facies, microfacies and gamma-ray spectrometry data: (A) drowning to periplatform turbidite setting; (B) drowning to (hemi)pelagic seamount setting and (C) drowning associated with the stratigraphical gap. In the lower Pa. hassi zone, rapid subsidence caused the platform to drown locally along the N–S to NW–SE trending faults (type A drowning). In the upper Pa. rhenana to the Pa. linguiformis zone, the drowning accelerated in the western part of the MSB due to locally higher subsidence rates combined with the Late Frasnian biotic crisis (type B). In the southern part of the basin, the platform emerged shortly before the Frasnian/Famennian (F/F) boundary and drowned in the Early to Late Famennian (type C). The primary cause of drowning was differential subsidence at the Laurussian passive margin. Eustatic sea-level fluctuations, if any, contributed only to a minor extent to the Late Frasnian drowning, but were effective in type C drowning during the Famennian. The drowning boundaries are associated with increased contents of K and Th, reflecting the deceleration of carbonate production. Uranium contents display isolated peaks that roughly correlate with the drowning boundaries or the stratigraphic gaps associated with the F/F boundary. The uranium contents are considered to reflect local depositional conditions and are not suitable for stratigraphic correlation. On the other hand, from the K and Th contents, we can infer Late Frasnian sea-level fluctuations with duration on the order of 1 Myr. These cyclic variations in K and Th contents proved to be useful in platform-to-basin stratigraphic correlation.     

Functional integration for enrolment constrains evolutionary variation of phacopid trilobites despite developmental modularity

Modularity and integration are variational properties expressed at various levels of the biological hierarchy. Mismatches among these levels, for example developmental modules that are integrated in a functional unit, could be informative of how evolutionary processes and trade‐offs have shaped organismal morphologies as well as clade diversification. In the present study, we explored the full, integrated and modular spaces of two developmental modules in phacopid trilobites, the cephalon and the pygidium, and highlight some differences among them. Such contrasts reveal firstly that evolutionary processes operating in the modular spaces are stronger in the cephalon, probably due to a complex regime of selection related to the numerous functions ensured by this module. Secondly, we demonstrate that the same pattern of covariation is shared among species, which also differentiate along this common functional integration. This common pattern might be the result of stabilizing selection acting on the enrolment and implying a coordinate variation between the cephalon and the pygidium in a certain direction of the morphospace. Finally, we noticed that Austerops legrandi differs slightly from other species in that its integration is partly restructured in the way the two modules interact. Such a divergence can result from the involvement of the cephalon in several vital functions that may have constrained the response of the features involved in enrolment and reorganized the covariation of the pygidium with the cephalon. Therefore, it is possible that important evolutionary trade‐offs between enrolment and other functions on the cephalon might have partly shaped the diversification of trilobites.     

Devonian Vertebrates From Colombia

Vertebrate remains are reported from the Emsian–Eifelian Floresta Formation and the Late Devonian (?Frasnian) Cuche Formation of north‐eastern Colombia. The material from the Floresta Formation is associated with a marine invertebrate fauna and includes an arthrodire and probably a rhenanid. Several vertebrate‐bearing localities are recorded from the Cuche Formation; vertebrates occur with plant remains and lingulid fragments. They include an acanthodian (Cheiracanthoides? sp.), a chondrichthyan (Antarctilamna? sp.), placoderms (Bothriolepis sp., Asterolepis? sp. and an undetermined groenlandaspidid or primitive brachythoracid arthrodire), a stegotrachelid actinopterygianand three sarcopterygians (a cosmine‐covered form tentatively referred to an osteolepidid, the porolepiform Holoptychiussp., and the rhizodontid Strepsodus? sp.). This assemblage suggests a Late Frasnian age and is surprisingly similar to Late Devonian vertebrate assemblages found in similar facies of Europe and North America, notwithstanding the presence of the Gondwanan chondrichthyan Antarctilamna?. key words: Vertebrata, Devonian, Colombia, South America, biostratigraphy, palaeobiogeography.     

Trilobites Asteropyginae dévoniens d’Iran

Ten species of trilobites belonging to 4 genera of the sub-family Asteropyginae DELO are described from the Devonian (Givetian-Frasnian) of Iran. Three of which are new: Neocalmonia lutensis nov. sp., Neocalmonia yazdii nov. sp. and Radiopyge sardarensis nov. sp. The study of this material have required the revision of Heliopyge sharudensis(PILLET, 1974) from the Givetian of the Eastern Alborz Mountains and of Neocalmonia quadricosta PILLET, 1969, type-species of the genus Neocalmonia, from the Frasnian of Afghanistan. The recognition of genera and species in common with the Givetian and the Frasnian of Afghanistan emphasizes the close relations between these two regions. The genera Bradocryphaeus, Heliopyge, Neocalmonia, Radiopyge, represented by related species, are likewise present in Iran, Afghanistan and Western Europe. Their presence confirms that these regions belong, during the Devonian, to the North Gondwanian domain.     

Contrasting patterns of disparity suggest differing constraints on the evolution of trilobite cephalic structures during the Cambrian ‘explosion’

Trilobites are an abundant group of Palaeozoic marine euarthropods that appear abruptly in the fossil record c. 521 million years ago. Quantifying the development of morphological variation (or ‘disparity’) through time in fossil groups like trilobites is critical in understanding evolutionary radiations such as the Cambrian ‘explosion’. Here, I use geometric morphometrics to quantify ‘cumulative disparity’ in functionally-important structures within the trilobite cephalon across their initial radiation during Cambrian Series 2. Overall cephalic disparity increased rapidly and attained a maximum within several million years. This pattern is dominated by the cephalic outline (in particular the genal spines), reflecting rapid, convergent expansion to the extremes of morphospace in a few early families. In contrast, removing the outline and focusing on structures such as the glabella and eye ridges (associated with feeding and vision, respectively) showed a more gradual increase in disparity, closer in line with taxonomic diversity and supporting the hypothesis of a relatively accurate trilobite fossil record. These contrasting patterns suggest that disparity in different structures was constrained in different ways, with extrinsic (ecological) factors probably having the major impact on overall disparity. It also implies that patterns of disparity in isolated substructures cannot necessarily be taken individually as representative of overall morphologies.     

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.     

Evidence for shallow‐water ‘Upper Kellwasser’ anoxia in the Frasnian–Famennian reefs of Alberta,Canada

The Frasnian–Famennian extinction witnessed the global devastation of both level‐bottom and reef communities in low latitudes. Marine extinctions in offshore level‐bottom communities are associated with two widespread, transgressive, anoxic ‘Kellwasser Events’ that support an anoxia–extinction link. Typical Kellwasser facies of bituminous limestones and shales are not obviously recorded in shallow‐water settings, and thus, it is unclear whether anoxia played a role in reef losses. We evaluate geochemical, petrographic and facies evidence for oxygen restriction from an extremely shallow‐water carbonate platform in Alberta. Sequence stratigraphy places the Frasnian–Famennian boundary at a sequence boundary that tops a laminated mudstone and interrupts carbonate platform deposition. Two transgressive pulses have been identified, one of which is associated with the second, major transgression of T‐R cycle IId of the Devonian eustatic sea‐level curve. Geochemical proxies indicate that these transgressions were accompanied by influx of dysoxic or anoxic waters. Organic carbon and U enrichment in the Frasnian, particularly just below the Frasnian–Famennian boundary, points to episodic dysoxic conditions that probably persisted into the basal Famennian and were coincidental with the global Upper Kellwasser Event. This study provides the first evidence for the smoking gun of an anoxia‐driven extinction in very shallow waters, implicating this potent killer in the demise of the Devonian reefs.     

Sedimentary and biofacies records in calciturbidites at the Devonian-Carboniferous boundary in Moravia (Moravian-Silesian Zone, Middle Europe)

Summary At the Devonian/Carboniferous boundary, major climatic and oceanographic changes influenced sedimentation on carbonate platforms and in peri-platfrom asreas. Three deep-water carbonate successions in Moravia, which were selected to represent different paleotectonic settings, have been studied with the aim of testing the influence of eustatic, climatic and tectonic controls on sedimentation and conodont paleoecology and taphonomy. On the slopes of the wide carbonate platforms of the Moravian Karst Development (Lesní lom and Grygov sections), an exemplary highstand shedding systems developed in the upper Famennian (expansa Zone), marked by a pronounced thickness of their respective calciturbidite successions and an abundance of shallow-water skeletal grains.Palamatolepis— andBispathodus-dominated conodont assemblages contain an admixture ofPolygnathus representing a transported, near-shore component. The eustatic sea-level fall in the praesulcata Zone and the lowstand conditions at the D/C boundary resulted in a decline of carbonate platform production and condensed deposition or nondeposition. In the Lesní lom section, a condensed sequence of turrbiditic calcarenites and shales (Middle praesulcata—lowermost sulcata Zone) was followed by lime mud calciturbidites (sulcata and duplicata Zones). In the conodont assemblages, the first event in the Lower praesulcata Zone was associated with the reduction of ‘mesopelagic’Palmatopic and a bloom of epipelagicPolygnathus communis. The second event in the Middle praesulcata Zone corresponds to the onset of polygnathidprotogranthodid biofacies, indicating a carbonate slope environment. In the Grygov section, a pronounced thickening and upward-coarsening succession of tubiditic calcilutites through calcarenites and intraclast breccias, with poor palmatolepid-bispathodid connodont assemblages (expansa Zone), indicates a progradation of the calciturbidite system associated with sea-level highstand. After a break in sedimentation, covering the interval from the Lower praseulcata to the base of Lower crenulata Zone, thick-bedded, fine-grained calciturbidites were deposited in the Lower crenulata Zone, and are associated with poor, mixed assemblages where siphonodellids and polygnathids predominate. At the isosticha-Upper crenulata/Lower typicus boundary, coasre grained, turbiditic calcarenites and breccias rich in clastic quartz grains and mixed conodont assemblages with reworked Frasnian and Famennian conodonts indicate a deep erosion of the source area, presumably due totectonic uplift (relative lowstand). In the Jesenec section, on the flanks of the volcanic seamount (the Drahany Development), a deep-water Upper Famennian condensed succession of calciturbidites and presumably winnowed pelagic limestones is marked by conodont assemblages of palmatolepid-bispathodid biofacies. More proximal calciturbidites with mixed deep-water and shallowwater conodonts prograde at the top of the Upper Famennian succession (Middle to Upper expansa Zone). A striking hiatus, covering the interval from the Early preaesulcata to the base of Lower crenulata Zone, resulted from extreme condensation and submarine bottom current erosion due to sea-level lowstand in the late Famennian and early Tournaisian. The renewed middle Tournaisian calciturbidite sedimentation with strong evidence of erosion at the source area indicates global eustatic rise and tectonic uplift of the Drahany Development seamounts (relative lowstand). The earlier occurrence of the uplift in the Jesenec area, relative to the Grygov section, shows the advance of tectonic processes over time in the Moravian-Silesian basin (orogenic polarity) as a consequence of Variscan orogenic movements.     

Devonian conodont biogeography: Quantitative analysis of provinciality

Cluster analysis of conodont faunas from each of 17 Lower‐lower Upper Devonian zones and subzones (data as reported by Klapper and Johnson, 1980) reveal changing patterns of provinciality.

Provinciality, expressed by a differentiation into western Laurussian and proto‐Tethyan biogeographic regions, is moderate in the lower Lochkovian but is low or absent in the upper Lochkovian‐lower Pragian. Provincialism returns in the Pragian and reaches its maximum development during the Emsian. Most Australian faunas are distinct from those of western Laurussia and the proto‐Tethys. Conodont faunas from suspect terranes of western North America display no unusual biogeographic affinities. Provincialism declines during the Eifelian and is only weakly developed in Givetian‐lower Frasnian faunas.

Changing global sea level during the Devonian may explain the development of Devonian conodont provinciality. As proposed by Klapper and Johnson (1980), low provinciality is associated with low stands of sea level. Endemic faunas develop in isolated epeiric seas during intermediate stages of sea level rise. High stands of sea level ultimately drown barriers to faunal exchange and prompt a return to low provinciality conditions.