Julien Denayer | Université de Liège (original) (raw)

Papers by Julien Denayer

Kölner Forum für Geologie und Paläontologie, 2011

The Becco locality (Liège province), belongs to the Theux tectonic window and represents a proxim... more The Becco locality (Liège province), belongs to the Theux tectonic window and represents a proximal, probably fluvial,
environment corresponding to a channel infill. We present here a preliminary report of the fossil assemblage discovered at the locality. The Becco site has yielded a diverse flora of early seed plants including Moresnetia zalesskyi, Dorinnotheca streeli and Condrusia sp. This assemblage, characteristic of the Belgian Famennian, highlights the diversity of early spermatophytes in the country. Becco has also delivered a rich vertebrate fauna with antiarch, groenlandaspid and phyllolepid placoderms, diplacanthiform acanthodians, as well as actinopterygians and various sarcopterygians. The fossiliferous assemblage of Becco resembles those of several Devonian tetrapod-bearing localities, including that of Strud in Belgium, and could therefore provide a favorable palaeoecological setting in the search for early tetrapods.

he IGCP 596–SDS Symposium is an international and multidisciplinary meeting aiming at a better un... more he IGCP 596–SDS Symposium is an international and multidisciplinary meeting aiming at a better understanding of the interactions existing between climate changes and biodiversity during the mid-Palaeozoic period (Devonian and Carboniferous). A time when terrestrial ecosystems experienced a biodiversity boom and oceanic ecosystems suffered from catastrophic extinctions of different magnitudes (e.g. Taghanic, Upper Kellwasser and Hangenberg events). This symposium brings together geochemists, geologists, palaeontologists and sedimentologists within the frame of the International Geoscience Programme (IGCP) Project 596 (climate change and biodiversity patterns in the mid-Palaeozoic) and the Subcommission on Devonian Stratigraphy (SDS) of the International Union of Geological Sciences.

Abstract – The Famennian (Upper Devonian, c. 372 to 359 Ma) strata of Belgium have recently recei... more Abstract – The Famennian (Upper Devonian, c. 372 to 359 Ma) strata of Belgium have recently received much attention after the discoveries of early tetrapod remains and outstandingly preserved continental arthropods. The Strud locality has yielded a diverse ﬂora and fauna including seed-plants,
tetrapods, various placoderm, actinopterygian, acanthodian and sarcopterygian ﬁshes, crustaceans (anostracans, notostracans, conchostracans and decapods) and a putative complete insect. This fossil assemblage is one of the oldest continental – probably fresh-water – ecosystems with a considerable vertebrate and invertebrate diversity. The study of the palaeoenvironment of the Strud locality is crucial
because it records one of the earliest and most important phases of tetrapod evolution that took place after their emergence but before their terrestrialization. It raises the question of environmental and ecological conditions for the Devonian aquatic ecosystem and the selection pressures occurring at the onset of tetrapod terrestrialization. The present study characterized the ﬂuvial facies of the Upper
Famennian sedimentary rocks of Strud and the surrounding areas. The exceptional preservation of arthropods and plants in the main fossiliferous layers is explained by rapid burial in the ﬁne-grained sediment of the quiet and conﬁned ﬂood plain environment. Newly investigated fossiliferous sections
in the Meuse–Samson area led to the description and correlation of key sections (Strud, Wierde and Jausse sections, complemented by the less continuous Haltinne, Huy and Coutisse sections). Moreover, the investigated sections allowed a reviewof the age of the fossiliferous horizon, which is nowdeﬁnitely considered to be Late Famennian in age.

Nature, 2012

After terrestrialization, the diversification of arthropods and vertebrates is thought to have oc... more After terrestrialization, the diversification of arthropods and vertebrates is thought to have occurred in two distinct phases, the first between the Silurian and the Frasnian stages (Late Devonian period) (425–385 million years (Myr) ago), and the second characterized by the emergence of numerous new major taxa, during the Late Carboniferous period (after 345 Myr ago). These two diversification periods bracket the depauperate vertebrate Romer’s gap (360-345 Myr ago) and arthropod gap (385–325 Myr ago), which could be due to preservational artefact. Although a
recent molecular dating has given an age of 390 Myr for the Holometabola, the record of hexapods during the Early–Middle Devonian (411.5–391 Myr ago, Pragian to Givetian stages) is exceptionally sparse and based on fragmentary remains, which hinders the timing of this diversification. Indeed, although Devonian Archaeognatha are problematic, the Pragian of Scotland has given some Collembola and the incomplete insect Rhyniognatha, with its diagnostic dicondylic, metapterygotan mandibles. The oldest, definitively winged insects are from the Serpukhovian stage (latest
Early Carboniferous period). Here we report the first complete Late Devonian insect, which was probably a terrestrial species. Its ‘orthopteroid’ mandibles are of an omnivorous type, clearly not modified for a solely carnivorous diet. This discovery narrows the 45-Myr gap in the fossil record of Hexapoda, and demonstrates further a first Devonian phase of diversification for the Hexapoda, as in vertebrates, and suggests that the Pterygota diversified before and during Romer’s gap.

An uppermost Famennian (Strunian) coral assemblage has been recovered in the middle part of the Y... more An uppermost Famennian (Strunian) coral assemblage has been recovered in the middle part of the Yılanlı Formation of the Istanbul Zone (Zonguldak and Bartın areas, NW Turkey). In the Bartın area, the studied fossiliferous interval corresponds to a c. 30 m-thick unit of bioclastic to peloidal wackestone to packstone grading to grainstone and including two stromatoporoid biostromes. In the Zonguldak area, 60 km westward, the bioclastic facies is dominant. The rugose corals are mainly solitary taxa belonging to the genera Campophyllum, Bounophyllum, Amplexocarinia, and ?Metriophyllum, and only one colonial genus occurs: Pseudoendophyllum. This fauna is similar to that documented in Europe. The campophyllids and dibunophyllids are the main component of the Uppermost Famennian assemblages in S Belgium, N France, W Germany, NW and S Poland. The endophyllids occur in S Poland, Novaya Zemlya, and in the Ural Mountains. The Istanbul Zone is supposed to be situated in the central part of the Palaeotethys Ocean, along the southern margin of Laurussia during the uppermost Devonian and Carboniferous. The rugose corals indicate some relationship with the eastern part of Laurussia, or that both areas were under a common marine influence at this time. The global Hangenberg event was not recognized in the Turkish localities, except considering the disappearance of the corals, occurring less than 19 m below the Devonian-Carboniferous boundary based on the foraminifers. There is no major facies change through the boundary and the first Carboniferous corals (small Uralinia and Caninophyllum) appear 6 m above the D-C boundary.

This article provides the first taxonomic description of Ivorian (Late Tournaisian, Early Carboni... more This article provides the first taxonomic description of Ivorian (Late Tournaisian, Early Carboniferous) rugose coral associations from north-western

In a case study of two wells from the Namur synclinorium, the black shale-dominated Chokier Forma... more In a case study of two wells from the Namur synclinorium, the black shale-dominated Chokier Formation was analyzed for petrography, mineralogy and organic geochemistry. Thin section petrography revealed a different facies assemblage for each well and a total of six microfacies types. Whole rock mineralogical data largely confirms a facies-dependent mineral composition. All different microfacies types of locality 1 (MFT-1, laminated silty mudstone; MFT-2, laminated mud-clast-rich mudstone; MFT-3, calcareous bioclast-rich mudstone) and locality 2 (MFT-4, lenticular mudstone; MFT-5, burrow-mottled mudstone; MFT-6, burrowed silty laminated mudstone) show ample evidence of a distal shelf environment that is sourced by currents, which may be linked to seasonal (monsoonal) transport of sediment from land to sea. Erosive bedload transport is an important mechanism of sedimentation whereas accumulation by settling from the water column can be excluded for the majority of investigated strata. Frequently, endobenthic activity and other processes after deposition, e.g. winnowing and reworking, caused strong fabric modifications. Various, but simple ichnofabrics prove at least temporary dysoxic conditions as confirmed by relatively low TS/TOC ratios. With regard to sedimentary features, organic geochemistry data hints to organic matter preservation due to rapid burial rather than intense anoxity. Silicification is a widespread diagenetic feature independent from facies and locality. It is most likely linked to a high supply rate of terrestrially dissolved silica as indicated by sEM observations, paleogeographic constraints and sedimentary features.

a b s t r a c t A significant part of the collection of mid-and Late Cretaceous (Cenomanian, Camp... more a b s t r a c t A significant part of the collection of mid-and Late Cretaceous (Cenomanian, Campanian and Maastrichtian) gastropods, bivalves and worms described and/or illustrated by de Ryckholt in his seminal work Mélanges paléontologiques between 1854 and 1862, has recently been retraced in the historical collections at Liège University. Of the original collection, more than 206 specimens, including 196 nominal types (lectotypes and genotypes), all considered lost, are now available. The genotypes of the gastropod genera Tudicula de Ryckholt, 1862a and Prosopostoma de Ryckholt, 1862a are photographically illustrated for the first time. Prosopostoma bucculans, from the Cenomanian Bernissart Formation (formerly 'Tourtia de Tournai'), is here chosen as the type species of the genus Prosopostoma, a possible stromboid. We also reassess the stratigraphic age of the type localities under the revised stratigraphic framework of Belgium and a detailed account on the research history of this material, in order to provide a thorough scientific background for future study of this formidable collection.

. Geological map of the study area after . Black points show the position of the mud mounds sensu... more . Geological map of the study area after . Black points show the position of the mud mounds sensu . White points show the main sample locations.

The origin of winged insects (Pterygota), one of the planet's most diverse lineages of organisms,... more The origin of winged insects (Pterygota), one of the planet's most diverse lineages of organisms, is assumed to lie in the Devonian, but as an extremely sparse fossil record impedes our understanding of their early diversification, any well-preserved insect fossil from this time would be particularly valuable 1 . Garrouste et al. 2 described an arthropod from Upper Devonian freshwater sediments from the Strud locality in Belgium (,360 million years ago) as an insect, Strudiella devonica. However, based on a thorough re-investigation of the specimen, we conclude that this interpretation is untenable. Hence we believe that, like several other Devonian arthropod remains 3-7 , Strudiella is not an insect, and utmost diligence should be taken in interpreting poorly preserved Devonian arthropods.

The microbial-sponge-bryozoan-coral bioherm of Kongul Yayla in the Central Taurides (South Turkey... more The microbial-sponge-bryozoan-coral bioherm of Kongul Yayla in the Central Taurides (South Turkey) contains a rich and diversified fauna. Sponges and rugose corals are of particular interest. The most common taxa are Siphonodendron irregulare, S. pauciradiale, S. cf. intermedium, Lithostrotion araneum, L. decipiens, L. maccoyanum, Axophyllum aff. pseudokirsopianum, Palaeosmilia multiseptata, P. murchisoni, Clisiophyllum aff. keyserlingi, Amygdalophyllum sp., Rotiphyllum cf. densum, Amplexocarinia aff. cravenensis, Soshkineophyllum? sp. and Espielia tauridensis sp. nov. newly described here. The tabulate corals are mostly micheliniids, syringoporids, cladochonids and auloporids. Heterocorals and chaetetids are also present. Siphonodendron pauciradiale and Lithostrotion maccoyanum are the guide taxa for the RC7β biozone and indicate an late Asbian age for the bioherm. Facies and coral assemblage argue for a South-European affinity of the Kongul Yayla reef and probably for the whole Anatolian terrane.

Kölner Forum für Geologie und Paläontologie, 2011

Nature, 2012

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The Belgian Frasnian was investigated for a long time and displays well dated and correlatable se... more The Belgian Frasnian was investigated for a long time and displays well dated and correlatable sections. They allow to document the evolution of the carbonate platform and buildups, and their decease relating to the Frasnian crises. The sequence stratigraphy of the Middle and Upper Frasnian was revised here and has allowed to establish that the development of the buildups was well correlated with the third-order sequences. During the Middle Frasian, the onset and vertical growth of the three levels of reefs ("Arche", "Boverie" and "Lion") correspond to the transgressive system tract (TST) of the sequences. During the high-stand (HST) and the fallingstage system tracts (FSST), their vertical growth decreased, and they evolved to progradant carbonate platforms, 1 -3 km wide and un to140 m high (including the biohermal core), in which bounstone are replaced by packstonegrainstone, then by shallow-water and intertidal mudstone (FSST). There is no evidence for the development of atolls rimmed by stromatoporoid-coral bareers, as it is usually suggested. The final emersion of these reefplatforms in the distal areas of the basin, and of the carbonate platform in the proximal areas, stopped the carbonate production until the following transgression-regression sequence. The last (third) middle Frasnian regression caused the end of the "Lion reef" and of the Middle Frasnian type reef. During the beginning of the transgression of the first ("Aisemont sequence") of the two recognized upper Frasnian sequences, was the first crisis affecting corals and stromatoporoids. Subsequently the carbonate production never recovered as previously. Reddish microbial mudmounds ("Petit-Mont" type buildups), grew during the TST and HST of this first sequence. They are smaller than the previous buildups, reaching up to 300 m wide and 80 m high. Their growth was dominantly vertical and there is no marked progradation during the HST. During the FSST of the Aisemont sequence, shallow-water mudstones and stromatolites developed on their top, then their emersion stropped their development. During the last Upper Frasnian sequence ("Lambermont sequence") the extension of anoxic-dysoxic facies prevented the development of large buildups, and only 1 to 2 metres-wide micro-mudmounds have so far been recorded.

The Hun Member of the Yvoir Formation represents the uppermost part of the Hastarian Substage (Lo... more The Hun Member of the Yvoir Formation represents the uppermost part of the Hastarian Substage (Lower Tournaisian, Lower Carboniferous) in the southern part of the Condroz sedimentation area of the Namur-Dinant Basin (S Belgium). This 12 m-thick member is composed of sandy bioclastic limestone alternating with shale layers and cherts; it is interpreted as the lowstand system tract of the third order sequence #3 of Hance et al. (2001). In the Chansin quarry (Bocq valley), this member has yielded an abundant association of silicified fauna showing an exquisite preservation. The association is dominated by rugose and tabulate corals along with brachiopods whereas bryozoans, gastropods and trilobites are only minor components. The rugose corals fauna is composed of solitary trochoid, ceratoid or cylindroid forms belonging to Amplexus coralloides, cf. Amplexizaphrentis sp., Bradyphyllum sp., Caninia cornucopiae, C. aff. cornucopiae, Caninophyllum patulum, "Lophophyllum" konincki, Proheterelasma omaliusi, Rotiphyllum sp., Saleelasma delepinei, Siphonophyllia cylindrica, Sychnoelasma konincki, Zaphrentites delanouei, and Zaphrentites sp. The good preservation of some specimens allows the observation of the calicular features that are rarely observed in time-equivalent fauna. Moreover, several corals show coiled or attached protocorallites. The tabulate corals are mainly small-sized (5 cm in diameter) micheliniid colonies, cladochonids and isolated corallites of Beaumontia. The massive colonies commonly show growth rings and talons. This coral association corresponds to a moderate diversified level-bottom community in which the external morphologies converge towards a single habitus. The brachiopod fauna, which is currently under study, is quite diverse but dominated by spire-bearers: athyridides (e.g. Lamellosathyris lamellosa, Coveenia sp.), spiriferides (Unispirifer sp., Tylothyris laminosa), and spiriferinides (e.g. Syringothyris sp.). It also includes some productides, strophomenides (Leptagonia gr. analoga), orthotetides (Shellwienella sp.), orthides (Rhipidomella michelini), rhynchonellides and terebratulides. The degree of disarticulation of the shells is particularly high, especially among spiriferides and spiriferinides, and it is clear that they have been disturbed and displaced from their living position, but the transport was probably over a short distance as indicated by the preservation of some delicate structures (e.g. mucronate cardinal extremities, flanges). Such accumulations might have been produced by storm events as suggest by the non-orientation of the corals preserved in chert layers.

The Lower Carboniferous macrofauna of the eastern Anti-Atlas, with exception of the ammonoids, is... more The Lower Carboniferous macrofauna of the eastern Anti-Atlas, with exception of the ammonoids, is poorly studied. Herein we present results from ongoing research of the rugose coral fauna of the eastern Tafilalt based on collections made during two short field campaigns in 2011 and 2013. Specimens have been collected from a series of surface outcrops in the Jebel Begaa, Hassi Nebech, Zrigat plain and Gara El Itima areas. Most corals are from carbonated horizons of the Merdani and Zrigat formations (late Tournaisian to late Viséan), fewer samples are from more detrital facies of the same formations. The oldest assemblage representing a latest Tournaisian age is found in conglomeratic layers west of the Jebel Begaa. This diverse assemblage consists mainly of small and medium-sized solitary forms belonging to the generaRylstonia, Sychnoelasma, Siphonophyllia, Cyathoclisia and Merlewoodia sp. Massive limestone blocks (scarp breccias?) from the bottom of Jebel Begaa contain Sychnoelasma sp. and Siphonodendron scaleberense. Higher up, in a supposed interval of green shales, large-sized siphonophyllids most likely indicate the late Viséan. However, the first late Viséan marker is Dibunophyllum bipartitum from the upper part of the Jebel. A relatively rich assemblage has been recovered from the limestones forming the Hassi Nebech ridge. It contains small and large-sized solitary and colonial corals (e.g. Amplexizaphrentis ssp., Rylstonia sguilmensis, Amygdalophyllum sp., Solenodendron horsfeldi). This assemblage is late Viséan in age, most likely Asbian. Zaphrentids and caniniids occur blow the limestones in greenishbrownish shales. A very diverse Asbian assemblage has been found in the Zrigat plain. Small solitary corals are common in shaly facies, and represented by e.g. Rylstonia benecompacta, Bradiphyllum sp., Cyathaxonia rushiana. Large solitary corals are abundant in carbonate and siliciclastic facies, e.g. Merlewoodia sp., Pseudozaphrentoides alloiteaui, Dibunophyllum bipartitum, Koninckophyllum cf. destitum, Pareynia sp., Axophyllum pseudokirsopianum. Colonial corals are diverse and fasciculate forms dominate; Siphonodendron irregulare, S. pauciradiale, S. martini, Solenodendron furcatum, and Lithostrotion decipiens. At the Gara El Itima, a late Viséan coral assemblage from the highest carbonate levels of the Zrigat Formation comprises a diverse range of forms including Zaphrentites sp., Rylstonia laxocolumnata, Caninia sp., Dibunophyllum bipartitum, Clisiophyllum garwoodi?, Diphyphyllum sp., and Palaecis sp. This assemblage lacks typical coral marker for the Brigantian, although Brigantian ammonoids have been found in this stratigraphic level.

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