First Triassic record of the beetle family Permocupedidae (Insecta: Coleoptera): a peculiar example of a Lazarus taxon

The first Mesozoic representative of the extinct archostematan beetle family Permocupedidae, Frankencupes ultimus, gen. et sp. nov., is described based on two isolated elytra from the Lower Anisian (Middle Triassic) Röt Formation of Lower Franconia, Germany. The new fossil occurrence extends the range of the family from the Lower Wuchiapingian (Upper Permian) up to the Anisian, and represents a fine example of a Lazarus taxon in the fossil record of beetles.     

Refinements in the Upper Permian to Lower Jurassic stratigraphy of Karakorum, Pakistan

New sampling on critical intervals of the uppermost Permian and Triassic successions of the Northern Karakorum Terrain in the Karakorum Range (Pakistan) has refined the stratigraphy. Two types of successions may be distinguished in the Karakorum Range: a carbonate platform succession, spanning the whole interval from Upper Permian to Upper Triassic, possibly with several gaps; and a basinal succession, deposited from the Middle Permian to Early Carnian (Late Triassic), when the carbonate platform prograded into the basin. With the approaching and later docking of the Karakorum Block against the Asian margin closing the Paleo-Tethys, a portion of Karakorum emerged while another part subsided as a fore-deep, receiving clastics from the emerging Cimmerian Range. Molassic sediments filled the basin, whilst shallow-water carbonates transgressed over the emerged carbonate platform sometime between the latest Triassic and the Pliensbachian (Early Jurassic), with Cimmerian deformation occurring to the north. The age control is provided by conodonts, with assemblages of late Wuchiapingian, Changhsingian, Induan (Griesbachian and Dienerian), late Olenekian, early Anisian, late Ladinian, and early Carnian ages, respectively. Some information on the section around the P/T boundary is provided by palynology and isotopic C¹³ values. The dating of the Norian/Rhaetian platform is provided by foraminifers.     

Anisian (middle triassic) buildups of the Northern Dolomites (Italy): The recovery of reef communities after the permian/triassic crisis

Summary After the end-Permian crisis and a global ‘reef gap’ in the early Triassic, reefs appeared again during the early Middle Triassic. Records of Anisian reefs are rare in the Tethys as well as in non-Tethyan regions. Most Anisian reefs are known from the western part of the Tethys but there are only very few studies focused on biota, facies types and the paleogeographical situation of these reefs. From the eastern part of the Tethys, Anisian reefs, reefal buildups or potential reef-building organisms have been reported from different regions of southern China. Most of the Anisian reefs known from western and central Europe as well as from southern China seem to be of middle and late Pelsonian age. The study area is situated in the northern Dolomites (South Tyrol, Italy) southeast of Bruneck (Brunico). It comprises the area between Olang (Valdaora) and Prags (Braies). The study is based on detailed investigations of the regional geology, stratigraphy and lithofacies (R. Zühlke, T. Bechst?dt) as well as on a comprehensive inventory of Anisian reef organisms (B. Senowbari-Daryan, E. Flügel). These data are used in the discussion of the controls on the recovery of reefs during the early Middle Triassic. Most late Anisian reef carbonates studied are represented by allochthonous talus reef blocks of cubicmeter size. Small biostromal autochthonous mounds are extremely rare (Piz da Peres). The reef mounds as well as most of the reef blocks occur within the middle to late Pelsonian Recoaro Formation. They were formed on the middle reaches of carbonate ramps in subtidal depths, slightly above the storm wave base with only moderate water energy. Most lithotypes observed in the reef blocks correspond to sponge and/or algal bafflestones. Low-growing sessile organisms (Olangocoelia (sponge, alga?), sphinctozoan sponges, bryozoans, soleno-poracean algae, corals) and encrusting epibionts (sponges, porostromate algae, cyanophycean crusts, foraminifera, worms, microproblematica) created low cm-sized biogenic structures (bioconstructions) which baffled and bound sediment. Organic framework was only of minor importance; it is restricted to theOlangocoelia lithotype. Framework porosity was small in these reef mounds. Submarine carbonate cements, therefore, are only of minor importance s compared with Permian or Ladinian reefs. The relatively high number of lithotypes encountered in the reef blocks indicates a high biofacies diversity. Regarding the relative frequency, the diverse biota consist in descending order ofOlangocoelia, sponges (sphinctozoans, inozoans, siliceous sponges), bryozoans, porostromate algae and worm tubes. The sphinctozoans are characterized by small, mostly incrusting forms. The numerical diversity (species richness) is low compared with late Permian or Ladinian and late Triassic sphinctozoan faunas occurring within reefs. Following the sponges, monospecific bryozoans (Reptonoditrypa cautica Sch?fer & Fois) are the most common organisms in the reef limestones. Porostromate algae were restricted to areas within the bioconstructions not inhabited by sponges. The low-diverse corals had no importance in the construction of an organic framework. Surprisingly, microbial crusts are rare or even lacking in the investigated Anisian bioconstructions. This is in contrast to late Permian and Ladinian as well as Carnian reefs which are characterized by the abundance of specific organic crusts. The same comes true for‘Tubiphytes’ which is a common constituent in Permian, Ladinian and Carnian reef carbonates but is very rare in the Anisian of the Olang Dolomites. Instead of‘Tubiphytes’ different kinds of worm tubes (spirorbid tubes, Mg-calcitic tubes and agglutinated tubes) were of importance as epifaunal elements. Macrobial encrustations consisting of characteristic successions of sponges, bryozoans, algae, worm tubes and microproblematica seem to be of greater quantitative importance than in Ladinian reefs. Destruction of organic skeletons (predominantly of bryozoans) by macroborers (cirripedia?) is a common feature. The Anisian reef organisms are distinctly different from late Permian and from most Ladinian reef-builders. No Permian Lazarus taxa have been found. New taxa: Sphinctozoan sponges—Celyphia? minima n.sp.,Thaumastocoelia dolomitica n. sp.,Deningeria tenuireticulata n. sp.,Deningeria crassireticulata n. sp.,Anisothalamia minima n.g. n.sp., Inozoan sponges-Meandrostia triassica n.sp. Microproblematica-Anisocellula fecunda n.g. n.sp., Porostromate alga-Brandneria dolomitica n.g. n.sp. Most of our data are in agreement with the model described byFois & Gaetani (1984) for the recovery of reef-building communities during the Ansian but the biotic diversity seems to be considerably higher than previously assumed. Anisian deposition and the formation of the reef mounds within the Pelsonian Recoaro Formation of the Dolomites were controlled by the combined effects of synsedimentary tectonics and eustatic changes in sea-level. During several time intervals, especially the early Anisian (northern and western Dolomites: tectonic uplift), the early Pelsonian (eastern Dolomites: drowning) and the late Illyrian (wide parts of the Dolomites: uplift and drowning), the sedimentation was predominantly controlled by regionally different tectonic subsidence rates. The amount of terrigenous clastic input associated with synsedimentary tectonics (tectonic uplift of hinterlands) had a major influence on carbonate deposition and reef development. The re-appearance of reef environments in the Olang Dolomites was controlled by a combination of regional and global factors (paleogeographic situation: development of carbonate ramps; decreasing subsidence of horst blocks; reduced terrigenous input; moderate rise in sea-level).     

HORSETAILS AND SEED FERNS FROM THE MIDDLE TRIASSIC (ANISIAN) LOCALITY KÜHWIESENKOPF (MONTE PRÀ DELLA VACCA), DOLOMITES, NORTHERN ITALY

Abstract:  Well-preserved floras from the Alpine Early–Middle Triassic are rare, and thus our understanding of the vegetation in this area during this period of time continues to be incomplete. As a result, every new find represents a significant piece of information that deserves thoughtful consideration. Anisian (Middle Triassic) sphenophytes and pteridosperms have recently been recovered from the Kühwiesenkopf locality (Monte Prà della Vacca) in northern Italy. The sphenophytes are represented by stem fragments, strobili and isolated sporangiophore heads of Equisetites , as well as by a few specimens of Neocalamites sp. and Echinostachys sp. The pteridosperms include abundant remains of the peltasperm foliage type Scytophyllum bergeri . A second Scytophyllum species in this flora, S. apoldense , is regarded as conspecific with S. bergeri based on epidermal anatomy; the two morphotypes are interpreted as sun and shade leaves of a single biological species. The seed-bearing disc Peltaspermum bornemannii sp. nov. probably represents the female reproductive structure of S. bergeri . Additional pteridosperm remains include foliage assignable to Sagenopteris sp. and Ptilozamites sp., in both cases perhaps the earliest records of these genera.     

Insect Leaf-Chewing Damage Tracks Herbivore Richness in Modern and Ancient Forests

The fossil record demonstrates that past climate changes and extinctions significantly affected the diversity of insect leaf-feeding damage, implying that the richness of damage types reflects that of the unsampled damage makers, and that the two are correlated through time. However, this relationship has not been quantified for living leaf-chewing insects, whose richness and mouthpart convergence have obscured their value for understanding past and present herbivore diversity. We hypothesized that the correlation of leaf-chewing damage types (DTs) and damage maker richness is directly observable in living forests. Using canopy access cranes at two lowland tropical rainforest sites in Panamá to survey 24 host-plant species, we found significant correlations between the numbers of leaf chewing insect species collected and the numbers of DTs observed to be made by the same species in feeding experiments, strongly supporting our hypothesis. Damage type richness was largely driven by insect species that make multiple DTs. Also, the rank-order abundances of DTs recorded at the Panamá sites and across a set of latest Cretaceous to middle Eocene fossil floras were highly correlated, indicating remarkable consistency of feeding-mode distributions through time. Most fossil and modern host-plant pairs displayed high similarity indices for their leaf-chewing DTs, but informative differences and trends in fossil damage composition became apparent when endophytic damage was included. Our results greatly expand the potential of insect-mediated leaf damage for interpreting insect herbivore richness and compositional heterogeneity from fossil floras and, equally promisingly, in living forests.     

Herbivore-induced plant volatiles mediate behavioral interactions between a leaf-chewing and a phloem-feeding herbivore

Plants respond adaptively to herbivore stress in order to maintain fitness. Upon herbivore attack, plants emit blends of volatile organic compounds (VOCs) that differ from those that are constitutively emitted. These defense responses are typically specific to the identity of the attacking herbivore and often linked to the herbivore's feeding guild (e.g. chewing, phloem-feeding). Herbivores use plant volatiles to locate suitable host plants and changes in volatile emissions can affect host-plant location. Therefore, herbivores from separate feeding guilds can interact indirectly through the modulation of plant responses. In this study we tested how damage by an herbivore from one feeding guild affected the host-plant choice of an herbivore from a separate feeding guild, and vice versa. A chewing herbivore, the Colorado potato beetle (Leptinotarsa decemlineata), and a phloem feeding herbivore, the green peach aphid (Myzus persicae), were assayed in olfactometers to assess behavioral responses to odors emitted by potato plants (Solanum tuberosum) that were damaged by herbivores from the other feeding guild. Leptinotarsa decemlineata oriented more frequently towards undamaged plants compared to M. persicae damaged plants. Surprisingly, M. persicae preferred plants that were damaged by L. decemlineata, although previous studies had shown that they perform worse on these plants. Distinct differences were detected in the volatile profiles of herbivore-damaged and undamaged plants. Leptinotarsa decemlineata induced stronger volatile emissions compared to undamaged control plants, while M. persicae tended to suppress volatile emissions. These herbivores demonstrate contrasting induction of plant volatiles and behavioral responses. Exploring the nature of co-occurring herbivores and how they perceive potential hosts can play a significant role in understanding the ecological functions and community dynamics of plant plasticity and interactions with a variety of herbivores.     

New fossil insects from the Anisian (Lower to Middle Muschelkalk) from the Central European Basin (Germany and The Netherlands)

The Palaeozoic–Mesozoic transition is characterized not only by the most massive Phanerozoic mass extinction at the end of the Permian period, but also its extensive aftermath and a prolonged period of major biotal recovery during the succeeding Middle to Late Triassic. Particularly, Anisian insect species from units of the Lower to Middle Muschelkalk from the Central European Basin are rare. The specimens described here originated from the Anisian Wellenkalk facies (Lower Muschelkalk), Vossenveld Formation of the Winterswijk quarry, The Netherlands, and from the orbicularis Member (lowermost Middle Muschelkalk, Anisian) of Esperstedt near Querfurt (Saxony-Anhalt). Thus, the described insect remains from Winterwijk and Esperstedt expand our knowledge about Middle Triassic terrestrial arthropod communities and their palaeodiversity. A new species of Chauliodites (C. esperstedti sp. nov) is introduced.     

New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

Proteromorphosis of Neospathodus (Conodonta) during the Permian–Triassic crisis and recovery

The Permian–Triassic evolution of platform conodonts (Gondolellidae) consists mainly of the development of the carina and the platform. During the sublethal environmental stress conditions subsequent to the Permian–Triassic extinction, the Wuchiapingian–Griesbachian Clarkina lineage is replaced by the primitive looking platform-lacking Dienerian–Aegean Neospathodus kummeli–Kashmirella timorensis lineage. It is assumed that, above Jinogondolella denticulata, end of the Capitanian Jinogondolella lineage, “Neospathodus” arcucristatus, an atavistic blade-like homeomorph that lacks a platform, underlies Protoclarkina crofti, of the base of the anagenetic Clarkina lineage. These primitive-looking forms are derived from their immediate ancestors by retrograde evolution, a phenomenon that has been described as proteromorphosis. Such events suggest that proteromorphosis occurs during periods of crisis, with the sudden reappearance of homeomorphic forms that are atavistic representatives of the clade. The phenomenon is further substantiated by several additional retrogradations that pace the Triassic, a period prone to such events.     

Permian predecessors of the mesozoic pteridosperms in western Angaraland,U.S.S.R.

The Kungurian and Upper Permian beds of western Angaraland yield a number of alleged predecessors of Mesozoic flora including Rhaphidopteris praecursoria sp. nov., the oldest representative of the genus hitherto known from the Jurassic only. R. praecursoria shows remarkably variable frond structures. Possible phyletic links between the Permian Rhaphidopteris, Triassic Corystospermaceae, and Jurassic Pachypteris and Rhaphidopteris are suggested.     

Callistophytalean pteridosperms from Permian aged floras of China

Abstract: Recent investigations into Permian aged floras from China have highlighted the widespread occurrence of callistophytalean pteridosperms that challenge previous understanding of their spatial and temporal distribution and diversity. In China, the group spans the Permian period and constitutes a distinctive but rare component in many peat‐forming environments. The stratigraphically earliest callistophytalean occurs in the Asselian‐Sakmarian stages with fossils from the Taiyuan Formation of northern China including ovules of Callospermarion undulatum in coal ball assemblages, and ovulate fronds of Norinosperma shanxiensis and synangiate fronds of Norinotheca shanxiensis in adpression assemblages. More abundant in the fossil records are adpression remains from the Roadian‐Wordian stages with the Lower Shihhotse Formation preserving abundant vegetative and ovulate remains of Emplectopteris triangularis that is now considered to represent a callistophytalean. The youngest callistophytalean recognised is from the Wuchaipingian‐Changhsingian stages with the Xuanwei Formation of southern China containing a single stem of Callistophyton boyssetii that provides indisputable evidence of the group in the lead up to the end‐Permian mass extinction. These accounts are augmented by analysis of pollen records that demonstrate the callistophytalean pollen genus Vesicaspora to be widespread through palynological assemblages from the Permian period in both North and South China, including the Upper Shihhotse Formation, Shihchienfeng Group, Xuanwei Formation, and possibly also in the mid‐Pennsylvanian Benxi Formation. Although macrofossil specimens are uncommon elements in the assemblages that contain them, they demonstrate the continuity of callistophytalean pteridosperms from the Pennsylvanian sub‐period into the early Guadalupian epoch of the Permian in North China and into the Lopingian epoch of the Permian in South China. Of the species present, both Callistophyton boyssetii and Callospermarion undulatum are known from the Pennsylvanian–earliest Permian age floras of Euramerica, whereas Norinosperma, Norinotheca and Emplectopteris appear to represent endemic Cathaysian elements. Results imply that callistophytalean pteridosperms can no longer be excluded from theories of post‐Carboniferous plant evolution and floristics, appearing to have played an important role in both Permian and Carboniferous aged plant communities. The presence of Vesicaspora in several formations from which macro‐remains have not been identified is a hopeful indicator that further callistophytalean pteridosperms are yet to be found.     

Anisian floras from the NE Iberian Peninsula and Balearic Islands: A synthesis

This paper focuses on the diversity and palaeontological significance of the Anisian floral record in the Triassic of the northeastern Iberian Peninsula. A summary of published information is followed by new records from macrofloral localities at Rodanas (Aragonese Branch of the Iberian Range) and at Maya del Baztán (Basque-Cantabrian Domain, Pyrenees). Both of these new localities also yielded three Anisian microfloral assemblages. These records provide new evidence of Anisian (basal Middle Triassic) floras in Spain. They are found to be somewhat similar to the classical flora of the Grès à Voltzia Formation (Upper Buntsandstein) of the northern Vosges in north-eastern France, of whose age has been shown to be early Anisian.     

The Permian–Triassic transition and the onset of Mesozoic sedimentation at the northwestern peri-Tethyan domain scale: Palaeogeographic maps and geodynamic implications

The main aim of this paper is to review Middle Permian through Middle Triassic continental successions in European. Secondly, areas of Middle–Late Permian sedimentation, the Permian–Triassic Boundary (PTB) and the onset of Triassic sedimentation at the scale of the westernmost peri-Tethyan domain are defined in order to construct palaeogeographic maps of the area and to discuss the impact of tectonics, climate and sediment supply on the preservation of continental sediment.At the scale of the western European peri-Tethyan basins, the Upper Permian is characterised by a general progradational pattern from playa-lake or floodplain to fluvial environments. In the northern Variscan Belt domain, areas of sedimentation were either isolated or connected to the large basin, which was occupied by the Zechstein Sea. In the southern Variscan Belt, during the Late Permian, either isolated endoreic basins occurred, with palaeocurrent directions indicating local sources, or basins underwent erosion and/or there was no deposition. These basins were not connected with the Tethys Ocean, which could be explained by a high border formed by Corsica–Sardinia palaeorelief and even parts of the Kabilia microplate. The palaeoflora and sedimentary environments suggest warm and semi-arid climatic conditions.At the scale of the whole study area, an unconformity (more or less angular) is observed almost everywhere between deposits of the Upper Permian and Triassic, except in the central part of the Germanic Basin. The sedimentation gap is more developed in the southern area where, in some basins, Upper Permian sediment does not occur. The large sedimentary supply, erosion and/or lack of deposition during the Late Permian, as well as the variable palaeocurrent direction pattern between the Middle–Late Permian and the Early Triassic indicate a period of relief rejuvenation during the Late Permian. During the Induan, all the intra-belt basins were under erosion and sediment was only preserved in the extra-belt domains (the northern and extreme southern domains). In the northern domain (the central part of the Germanic Basin), sediment was preserved under the same climatic conditions as during the latest Permian, whereas in the extreme southern domain, it was probably preserved in the Tethys Ocean, implying a large amount of detrital components entering the marine waters. Mesozoic sedimentation began in the early Olenekian; the ephemeral fluvial systems indicate arid climatic conditions during this period. Three distinct areas of sedimentation occur: a northern and southern domain, separated by an intra-belt domain. The latter accumulated sediments during the Early–Middle Permian and experienced erosion and/or no-deposition conditions between the Middle–Late Permian and the beginning of Mesozoic sedimentation, dated as Anisian to Hettangian. At the top of the Lower Triassic, another tectonically induced, more or less angular unconformity is observed: the Hardegsen unconformity, which is dated as intra-Spathian and is especially found in the North European basins. This tectonic activity created new source areas and a new fluvial style, with marine influences at the distal part of the systems. During the Anisian and Ladinian, continental sedimentation was characterised by a retrogradational trend. In other words, the fluvial system evolved into fluvio-marine environments, attesting to a direct influence of the Tethys Ocean in the southern and northern domains. Both at the end of the Olenekian (Spathian) and during the Anisian, the presence of palaeosols, micro- and macrofloras indicate less arid conditions throughout this domain.     

Facies and biota of Anisian to Carnian carbonate platforms in the Northern Calcareous Alps (Tyrol and Bavaria)

Summary During the Middle and early Late Triassic carbonate ramps and rimmed platforms developed at the northwestern margin of the Tethys ocean. In the Northern Calcareous Alps, Anisian stacked homoclinal ramps evolved through a transitional stage with distally steepened ramps to huge rimmed platforms of Late Ladinian to Early Carnian age. Middle Triassic to early Late Triassic facies and biota of basin, slope and platform depositional systems are described. Special emphasis is given to foraminifers, sponges, microproblematic organisms and algae. The Ladinian to early Carnian reef associations are characterized by the abundance of segmented sponges, microproblematica, biogenic crusts and synsedimentary cements. Among the foraminifers, recifal forms likeHydrania dulloi andCucurbita infundibuliformis (Carnian in age) are reported from the Northern Calcareous Alps for the first time. Some sphinctozoid sponges likeParavesicocaulis concentricus were known until now only from the Hungarian and Russian Triassic.     

Biostratigraphy and palaeobiogeography of Lower Permian (lower Kungurian) conodonts from the Tak Fa Formation (Saraburi Limestone), Thailand

Lower Permian (lower Kungurian) conodonts are reported from the Indochina Block of Southeast Asia. The fauna from the Tak Fa Formation of the Saraburi Limestone Group exposed in limestone hills NNW of Khok Samrong, Thailand, includes Sweetognathus subsymmetricus Wang, Ritter and Clark (early forms) and Pseudosweetognathus costatus Wang, Ritter and Clark. The co-occurrence of these species indicates a Lower Permian age (upper half of the lower Kungurian) for the sampled limestones representative of the Mesogondolella siciliensis–S. subsymmetricus Zone of South China. Sweetognathus and Pseudosweetognathus are for the first time reported from the Indochina terrane located in the palaeoequatorial belt in the Kungurian. Pseudosweetognathus appears restricted to the Kungurian of the South China and Indochina terranes thus supporting palaeogeographic reconstructions that isolate these terranes within the Palaeo-Tethys during the Kungurian.     

Re-evaluation of Halle’s fertile pteridosperms from the Permian floras of Shanxi Province, China

In 1927, T. G. Halle published an extensive synthesis of the Permian fossil plant assemblages from China’s Shanxi Province that included five genera and species of seed plant that either had ovules in organic attachment or closely and repeatedly associated with foliage. These fertile pteridosperms were the subject of a later publication (1929) that provided additional information on each species, but since that time only Nystroemia pectiniformis Halle has been subjected to detailed investigation based on the original materials. Reinvestigation of the remaining four species has yielded additional examples and new information for each species, and has confirmed the pteridospermalean affinity for three of the four species. Alethopteris norinii Halle shows organic attachment of a pollen organ similar to Wittleseya (Newberry) Halle and is reinterpreted as a microsporangiate medullosan pteridosperm. Emplectopteris triangularis Halle and Sphenopteris tenuis (Schenk) Halle both bear numerous ovules attached directly to the leaf laminae, with Emplectopteris conforming with the gross-organisation of callistophytalean pteridosperms and S. tenuis being interpreted as a derived lagenostomalean pteridosperm. However, discovery of the counterpart to the key specimen of Pecopteris wongii Halle that Halle interpreted as bearing ovules demonstrates this species to be a sterile marattialean frond that agrees with its assignment to Pecopteris. In addition, two further specimens are demonstrated to be fertile organs of previously unknown pteridosperms; Norinosperma shanxiensis gen. et sp. nov. bears cardiocarpalean ovules abaxially on the proximal portions of the frond with ovules positioned directly on the veins of a leaf lamina; Norinotheca shanxiensis gen. et sp. nov. bears large pollen organs abaxially and singly on the vein of the leaf lamina. Collectively Halle’s pteridosperms remain important today as they are some of the few specimens that actually demonstrate organic connection of fertile organs to foliage, helping to characterise the morphology and organisation of this seed plant group. These species also demonstrate the persistence of seed plant families and genera in the Permian of China that became regionally extinct at the end of the Carboniferous in Euramerica, and show that pteridosperms remained an important component of Permian as well as Carboniferous floras globally.     

A sequence stratigraphic approach to a Middle Triassic shelf-slope complex of the Ligurian Alps (Ligurian Briançonnais,Monte Carmo-Rialto unit,Italy)

The Mesozoic sedimentary cover belonging to the Monte Carmo-Rialto unit of the Ligurian Briançonnais domain is composed of Scythian clastics and Anisian to Carnian carbonate rocks over 300 m thick. This paper focuses on the stratigraphy of this carbonate complex, its environmental significance, and its evolution in light of dynamic stratigraphy. Our facies analysis of limestones and dolomites of the Triassic complex allowed us to reconstruct an environmental model. Data support a distally steepened carbonate ramp of Anisian age evolving to a more diversified Ladinian platform with an oolitic sand-bar belt separating the lagoon from the slope. The Monte Carmo-Rialto slope facies are the only witnesses of deep sedimentation in the Triassic terrains of the Ligurian Briançonnais domain, otherwise represented by shallow-water carbonate deposits. On the basis of facies succession, we have identified nine medium-scale cycles (third-order sequences) in the study area, comparable to those evidenced in the Briançonnais s.s. domain by the French authors. Small-scale cycles analysis evidenced mainly shallowing-upward trends in the examined sequences; although a few evidences of transgression-related deposits (deepening upward cycles) have been found at the base three sequences, they have been mostly obliterated by dolomitization and masked by local tectonics. For this reason, we can undoubtedly distinguish only the part of each sequence belonging to HST, while the TST, though present, still remains a partition that cannot be precisely characterized. In the same way, LSTs are not present in the Monte Carmo-Rialto unit, due to the original relative landward position of the examined area. Sequence stratigraphy analysis indicates different long-term dynamics for the two evolutionary stages of the Triassic Ligurian platform: a general landward backstepping to moderate progradation during the Early Anisian and true progradation during the latest Anisian and Ladinian. In addition, a good fit with the sequences proposed by the SEPM chart has been found, indicating a correspondence for the third-order sequences of the Middle Triassic.     

Herbivore-herbivore interactions complicate links between soil fertility and pest resistance

Soil fertility is tightly linked with herbivore pressure because it affects the nutritional status of host plants as well as the production of anti-herbivore defenses. This in turn can influence whether herbivores in different feeding guilds render plants more or less susceptible to one another. Thus, growers’ fertility management choices may impact herbivores through a variety of indirect channels. We examined relationships between soil fertility and interactions between phloem-feeding and leaf-chewing herbivores on broccoli (Brassica oleracea) plants in the greenhouse, taking advantage of natural variation in nitrogen (N) and phosphorus (P) in soils from 20 working organic vegetable farms. Next, we experimentally fertilized soil in a field trial with N and/or P to examine the consequences of these nutrients for growth of and interactions between specialist and generalist herbivores. Soils on our cooperating farms varied widely in P and N concentrations, with 40% exceeding recommended pre-plant N concentrations and 90% exceeding P recommendations. In single-herbivore infestations, augmenting N in the soil increased caterpillar (Pieris rapae) growth, augmented N and P additively enhanced generalist green peach aphid (Myzus persicae) colonization, and augmented P (but not N) increased specialist cabbage aphid (Brevicoryne brassicae) growth. In dual-guild herbivore infestations, caterpillars facilitated specialist cabbage aphid growth in the absence of fertilizer, but this pattern disappeared under augmented N, and reversed under augmented P. We found that a complex web of indirect effects linked soil fertility to herbivore performance, depending on the identity of the nutrients being altered, the ecological roles of responding herbivore species (i.e., specialist versus generalist), and indirect interactions between chewing and sucking herbivores. More generally, we highlight that successful use of fertility management to improve pest resistance requires careful consideration of herbivore feeding niches and herbivore-herbivore interactions.     

A Middle Triassic thoracopterid from China highlights the evolutionary origin of overwater gliding in early ray-finned fishes

Gliding adaptations in thoracopterid flying fishes represent a remarkable case of convergent evolution of overwater gliding strategy with modern exocoetid flying fishes, but the evolutionary origin of this strategy was poorly known in the thoracopterids because of lack of transitional forms. Until recently, all thoracopterids, from the Late Triassic of Austria and Italy and the Middle Triassic of South China, were highly specialized ‘four-winged’ gliders in having wing-like paired fins and an asymmetrical caudal fin with the lower caudal lobe notably larger than the upper lobe. Here, we show that the new genus Wushaichthys and the previously alleged ‘peltopleurid’ Peripeltopleurus, from the Middle Triassic (Ladinian, 235–242 Ma) of South China and near the Ladinian/Anisian boundary of southern Switzerland and northern Italy, respectively, represent the most primitive and oldest known thoracopterids. Wushaichthys, the most basal thoracopterid, shows certain derived features of this group in the skull. Peripeltopleurus shows a condition intermediate between Wushaichthys and Thoracopterus in having a slightly asymmetrical caudal fin but still lacking wing-like paired fins. Phylogenetic studies suggest that the evolution of overwater gliding of thoracopterids was gradual in nature; a four-stage adaption following the ‘cranial specialization–asymmetrical caudal fin–enlarged paired fins–scale reduction’ sequence has been recognized in thoracopterid evolution. Moreover, Wushaichthys and Peripeltopleurus bear hooklets on the anal fin of supposed males, resembling those of modern viviparious teleosts. Early thoracopterids probably had evolved a live-bearing reproductive strategy.