The early Eocene birds of the Messel fossil site: a 48 million‐year‐old bird community adds a temporal perspective to the evolution of tropical avifaunas

Birds play an important role in studies addressing the diversity and species richness of tropical ecosystems, but because of the poor avian fossil record in all extant tropical regions, a temporal perspective is mainly provided by divergence dates derived from calibrated molecular analyses. Tropical ecosystems were, however, widespread in the Northern Hemisphere during the early Cenozoic, and the early Eocene German fossil site Messel in particular has yielded a rich avian fossil record. The Messel avifauna is characterized by a considerable number of flightless birds, as well as a high diversity of aerial insectivores and the absence of large arboreal birds. With about 70 currently known species in 42 named genus‐level and at least 39 family‐level taxa, it approaches extant tropical biotas in terms of species richness and taxonomic diversity. With regard to its taxonomic composition and presumed ecological characteristics, the Messel avifauna is more similar to the Neotropics, Madagascar, and New Guinea than to tropical forests in continental Africa and Asia. Because the former regions were geographically isolated during most of the Cenozoic, their characteristics may be due to the absence of biotic factors, especially those related to the diversification of placental mammals, which impacted tropical avifaunas in Africa and Asia. The crown groups of most avian taxa that already existed in early Eocene forests are species‐poor. This does not support the hypothesis that the antiquity of tropical ecosystems is key to the diversity of tropical avifaunas, and suggests that high diversification rates may be of greater significance.     

The importance of Planolites in the Cambrian substrate revolution

Early Cambrian subtidal shelf substrates were characterized by low water content and steep chemical gradients, conditions likely facilitated by the presence of microbial mats as reflected by an abundance of microbially-mediated sedimentary structures in Lower Cambrian strata. Such substrate conditions would have been unfavourable for burrowing by benthic metazoans. A combination of environmental restrictions and a lack of adaptations to vertical burrowing likely prevented most benthic metazoans from burrowing infaunally in Early Cambrian subtidal shelf substrates. The eventual acquisition of burrowing adaptations by benthic metazoans later in the Cambrian promoted an increase in the depth and intensity of bioturbation and initiated a transition toward well-hydrated substrates in which extensive infaunal activity was possible.Siliciclastic units of the Lower Cambrian succession in the White–Inyo Mountains, eastern California, contain abundant horizontal bioturbation on bedding planes, as documented by bedding plane bioturbation indices, but little vertical bioturbation, as shown by ichnofabric indices and x-radiography. Planolites, a simple horizontal trace fossil, represents the dominant type of bioturbation in these units. Planolites is found in a range of diameters, indicating that more than one species of tracemaker likely produced this type of trace. Although these Planolites do not have a vertical component, their abundance on bedding planes indicates that the activities of Planolites tracemakers had a significant impact on subtidal shelf substrates, represented by Lower Cambrian units in the White–Inyo Mountains, early in the Cambrian substrate revolution.     

Origin and dispersal of the earliest brachiopods

Brachiopods first appeared at the very beginning of the Phanerozoic together with the first skeletal organisms. Most brachiopod taxa that arose in the first half of the Cambrian had a short temporal range and became completely extinct by the middle of the Middle Cambrian. Rigid articulation of the valves of brachiopods was provided by various structures, which also appeared in the Early Cambrian. This fact points to the importance of this feature for the formation of the whole group and at the same time testifies to the high variability of rigid articulation at the early stages of brachiopod evolution. This is a typical manifestation of archaic diversity in this animal phylum, which appeared very early in the Phanerozoic. Another important property of the archaic diversity of the early brachiopods was the large number of centers of diversification. As for the majority of groups, climatic zonality was the main factor determining the distribution of brachiopods at the beginning of the Phanerozoic. The main ecological types of brachiopods also appeared in the Early Cambrian.     

Branching archaeocyaths as ecosystem engineers during the Cambrian radiation

The rapid origination and diversification of major animal body plans during the early Cambrian coincide with the rise of Earth's first animal-built framework reefs. Given the importance of scleractinian coral reefs as ecological facilitators in modern oceans, we investigate the impact of archaeocyathan (Class Archaeocyatha) reefs as engineered ecosystems during the Cambrian radiation. In this study, we present the first high-resolution, three-dimensional (3D) reconstructions of branching archaeocyathide (Order Archaeocyathida) individuals from three localities on the Laurentian paleocontinent. Because branched forms in sponges and corals display phenotypic plasticity that preserve the characteristics of the surrounding growth environment, we compare morphological measurements from our fossil specimens to those of modern corals to infer the surface conditions of Earth's first reefs. These data demonstrate that archaeocyaths could withstand and influence the flow of water, accommodate photosymbionts, and build topographically complex and stable structures much like corals today. We also recognize a stepwise increase in the roughness of reef environments in the lower Cambrian, which would have laid a foundation for more abundant and diverse coevolving fauna.     

Exceptionally preserved jellyfishes from the Middle Cambrian

Cnidarians represent an early diverging animal group and thus insight into their origin and diversification is key to understanding metazoan evolution. Further, cnidarian jellyfish comprise an important component of modern marine planktonic ecosystems. Here we report on exceptionally preserved cnidarian jellyfish fossils from the Middle Cambrian (approximately 505 million years old) Marjum Formation of Utah. These are the first described Cambrian jellyfish fossils to display exquisite preservation of soft part anatomy including detailed features of structures interpreted as trailing tentacles and subumbrellar and exumbrellar surfaces. If the interpretation of these preserved characters is correct, their presence is diagnostic of modern jellyfish taxa. These new discoveries may provide insight into the scope of cnidarian diversity shortly after the Cambrian radiation, and would reinforce the notion that important taxonomic components of the modern planktonic realm were in place by the Cambrian period.     

How far did feedback between biodiversity and early diagenesis affect the nature of Early Palaeozoic sea floors?

Latest Precambrian to Early Palaeozoic biosphere evolution triggered changes in early diagenesis and carbonate precipitation which fed back to biodiversity through colonization of hard substrates. Progressive increase in the depth and intensity of bioturbation and bio‐irrigation lowered the zone of early carbonate cementation in the uppermost sediment column. This firstly led to a decline in the abundance of the flat‐pebble conglomerates which had been a common feature of Cambrian and Early Ordovician successions, replaced by the peak and subsequent decline in the Palaeozoic abundance of submarine hardgrounds. The availability of very widespread lithified sea floors in shallow subtidal settings during the Ordovician promoted a rapid expansion in sclerobiont diversity and contributed to the Great Ordovician Biodiversification Event.     

The microbial landscape in bioturbated mangrove sediment: A resource for promoting nature-based solutions for mangroves

Globally, soils and sediments are affected by the bioturbation activities of benthic species. The consequences of these activities are particularly impactful in intertidal sediment, which is generally anoxic and nutrient-poor. Mangrove intertidal sediments are of particular interest because, as the most productive forests and one of the most important stores of blue carbon, they provide global-scale ecosystem services. The mangrove sediment microbiome is fundamental for ecosystem functioning, influencing the efficiency of nutrient cycling and the abundance and distribution of key biological elements. Redox reactions in bioturbated sediment can be extremely complex, with one reaction creating a cascade effect on the succession of respiration pathways. This facilitates the overlap of different respiratory metabolisms important in the element cycles of the mangrove sediment, including carbon, nitrogen, sulphur and iron cycles, among others. Considering that all ecological functions and services provided by mangrove environments involve microorganisms, this work reviews the microbial roles in nutrient cycling in relation to bioturbation by animals and plants, the main mangrove ecosystem engineers. We highlight the diversity of bioturbating organisms and explore the diversity, dynamics and functions of the sediment microbiome, considering both the impacts of bioturbation. Finally, we review the growing evidence that bioturbation, through altering the sediment microbiome and environment, determining a ‘halo effect’, can ameliorate conditions for plant growth, highlighting the potential of the mangrove microbiome as a nature-based solution to sustain mangrove development and support the role of this ecosystem to deliver essential ecological services.     

Palaeoceanographic and palaeoclimatic inferences from Late Cretaceous planktonic foraminiferal assemblages from the Exmouth Plateau (ODP Sites 762 and 763, eastern Indian Ocean)

The evolution of planktonic foraminifera during the Late Cretaceous is marked in the Santonian by the disappearance of complex morphotypes (the marginotruncanids), and the contemporary increasing importance and diversification of another group of complex taxa, the globotruncanids. Upper Turonian to lower Campanian planktonic foraminiferal assemblages from Holes 762C and 763B (Ocean Drilling Program, Leg 122, Exmouth Plateau, 47°S palaeolatitude) were studied in detail to evaluate the compositional variations at the genus and species level based on the assumption that, in the Cretaceous oceans as in the modern, any faunal change was associated with changes in the characteristics and the degree of stability of the oceanic surface waters. Three major groups were recognised based on gross morphology, and following the assumption that Cretaceous planktonic foraminifera, although extinct, had life-history strategies comparable to those of modern planktonics: 1 – r-selected opportunists; 2 – k-selected specialists; 3 – r/k intermediate morphotypes which include all genera that display a range of trophic strategies in-between opportunist and specialist taxa. Although planktonic foraminiferal assemblages are characterised by a progressive appearance of complex taxa, this trend is discontinuous. Variation in number of species and specimens within genera has allowed recognition of five discrete intervals each of them reflecting different oceanic conditions based on fluctuations in diversity and abundance of the major morphotypes. Planktonic forms show cyclical fluctuations in diversity and abundance of cold (r-strategists) and warm taxa (k-strategists), perhaps representing alternating phases of unstable conditions (suggesting a weakly stratified upper water column in a mesotrophic environment), and well-stratified surface and near-surface waters (indicating a more oligotrophic environment). Interval 1, middle Turonian to early Coniacian in age, is dominated by the r/k intermediate morphotypes which alternate with r-strategists. These cyclical alternations are used to identify three additional sub-intervals. Interval 2, aged middle to late Coniacian, is characterised by the increasing number of species and relative abundance of k-strategists. After this maximum diversification the k-strategists show a progressive decrease reaching a minimum value in Interval 3 (early to late Santonian), which corresponds to the extinction of the genus Marginotruncana. In the Interval 4, latest Santonian in age, the k-strategists, represented mainly by the genera Globotruncana, increase again in diversity and abundance. The last Interval 5 (early Campanian) is dominated by juvenile globotruncanids and r-strategists which fluctuate in opposite phase. The positive peak (Interval 2) related to the maximum diversification of warm taxa (k-strategists) in the Coniacian seems to correspond to a warmer episode. It is followed by a marked decrease in the relative abundance of warm taxa (k-strategists crisis) with a minimum in the late Santonian (Interval 3), reflecting a decrease in temperature. Detailed analysis of faunal variations allows the Santonian faunal turnover to be ascribed to a cooling event strong enough to cause the extinction of the marginotruncanids.     

A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in‐field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in‐field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes.     

Eco‐evolutionary feedbacks between private and public goods: evidence from toxic algal blooms

The importance of ‘eco‐evolutionary feedbacks’ in natural systems is currently unclear. Here, we advance a general hypothesis for a particular class of eco‐evolutionary feedbacks with potentially large, long‐lasting impacts in complex ecosystems. These eco‐evolutionary feedbacks involve traits that mediate important interactions with abiotic and biotic features of the environment and a self‐driven reversal of selection as the ecological impact of the trait varies between private (small scale) and public (large scale). Toxic algal blooms may involve such eco‐evolutionary feedbacks due to the emergence of public goods. We review evidence that toxin production by microalgae may yield ‘privatised’ benefits for individual cells or colonies under pre‐ and early‐bloom conditions; however, the large‐scale, ecosystem‐level effects of toxicity associated with bloom states yield benefits that are necessarily ‘public’. Theory predicts that the replacement of private with public goods may reverse selection for toxicity in the absence of higher level selection. Indeed, blooms often harbor significant genetic and functional diversity: bloom populations may undergo genetic differentiation over a scale of days, and even genetically similar lineages may vary widely in toxic potential. Intriguingly, these observations find parallels in terrestrial communities, suggesting that toxic blooms may serve as useful models for eco‐evolutionary dynamics in nature. Eco‐evolutionary feedbacks involving the emergence of a public good may shed new light on the potential for interactions between ecology and evolution to influence the structure and function of entire ecosystems.     

Butterflies of the Cambrian benthos? Shield position in bradoriid arthropods

Mode of preservation and method of recovery strongly influences our understanding of the life habits of extinct organisms. Bradoriid arthropods were abundant, and diverse members of early Cambrian ecosystems and most life reconstructions display these animals with the two shields of the carapace open in a ‘butterfly’ configuration. This favoured reconstruction is largely based on the abundance of ‘crack‐out’ specimens preserved in this position (e.g. Kunmingella from the early Cambrian of China). In contrast, large collections of acid processed bradoriids from the Arrowie Basin of South Australia (Cambrian Stage 3) are preserved with a narrow gape at the ventral margin or completely closed with the carapace folded along the dorsal midline. The relative abundance of conjoined, closed (or partially closed) specimens from the lower Cambrian Hawker Group succession suggests that at least some bradoriid taxa were capable of withdrawing appendages and tightly closing the shields, challenging the common view that the majority of bradoriids usually held their carapaces open in a ‘butterfly’ configuration during life. New data show that layers of the bradoriid carapace are continuous through the dorsal fold with no evidence for complex articulating structures as in ostracod hinges. The relatively pliable, sclerotized or lightly mineralized calcium phosphate composition of the carapace and the simple, flexible dorsal fold facilitated opening and closing of the shields. Despite not being closely related, ostracods share close biomechanical and ecological similarities with bradoriids. The evolution of more complex articulating hinge structures – together with well‐developed musculature – in ostracods during the Early Ordovician, may have provided more efficient means for shield articulation and movement, thus promoting the ecological success of ostracods throughout the Phanerozoic.     

The Ordovician Radiation: A Follow-up to the Cambrian Explosion?

There was a major diversification known as the Ordovician Radiation,in the period immediately following the Cambrian. This eventis unique in taxonomic, ecologic and biogeographic aspects. While all of the phyla but one were established during the Cambrianexplosion, taxonomic increases during the Ordovician were manifestat lower taxonomic levels although ordinal level diversity doubled.Marine family diversity tripled and within clade diversity increasesoccurred at the genus and species levels. The Ordovician radiationestablished the Paleozoic Evolutionary Fauna; those taxa whichdominated the marine realm for the next 250 million years. Communitystructure dramatically increased in complexity. New communitieswere established and there were fundamental shifts in dominanceand abundance. Over the past ten years, there has been an effort to examinethis radiation at different scales. In comparison with the Cambrianexplosion which appears to be more globally mediated, localand regional studies of Ordovician faunas reveal sharp transitionswith timing and magnitudes that vary geographically. These transitionssuggest a more episodic and complex history than that revealedthrough synoptic global studies alone. Despite its apparent uniqueness, we cannot exclude the possibilitythat the Ordovician radiation was an extension of Cambrian diversitydynamics. That is, the Ordovician radiation may have been anevent independent of the Cambrian radiation and thus requiringa different set of explanations, or it may have been the inevitablefollow-up to the Cambrian radiation. Future studies should focuson resolving this issue.     

Cobble community DNA as a tool to monitor patterns of biodiversity within kelp forest ecosystems

Kelp forest ecosystems dominate 150,000 km of global temperate coastline, rivalling the coastal occurrence of coral reefs. Despite the astounding biological diversity and productive ecological communities associated with kelp forests, patterns of species richness and composition are difficult to monitor and compare. Crustose coralline algae are a critically important substrate for propagule settlement for a range of kelp forest species. Coralline‐covered cobbles are home to hundreds of species of benthic animals and algae and form a replicable unit for ecological assays. Here, we use DNA metabarcoding of bulk DNA extracts sampled from cobbles to explore patterns of species diversity in kelp forests of the central California coast. The data from 97 cobbles within kelp forest ecosystems at three sites in Central California show the presence of 752 molecular operational taxonomic units (MOTUs) and 53 MOTUs assigned up to the species level with >95% similarity to current databases. We are able to detect spatial patterns of important management targets such as abalone recruits, and localized abundance of sea stars in 2012. Comparison of classic ecological surveys of these sites reveals large differences in species targets for these two approaches. In order to make such comparisons more quantitative, we use Presence/Absence Metabarcoding, using the fraction of replicate cobbles showing a species as a measure of its local abundance. This approach provides a fast and repeatable survey method that can be applied for biodiversity assessments across systems to shed light on the impact of different ecological disturbances and the role played by marine protected areas.     

Pacific walrus: Benthic bioturbator of Beringia

The dependency of walruses on sea ice as habitat, the extent of their feeding, their benthic bioturbation and consequent nutrient flux suggest that walruses play a major ecological role in Beringia. This suggestion is supported by several lines of evidence, accumulated during more than three decades of enquiry and leading to the hypothesis that positive feedbacks of walrus feeding strongly influence productivity and ecological function via benthic bioturbation and nutrient flux. Walruses annually consume an estimated 3 million metric tons of benthic biomass. Walrus prey species inhabit patches across the shelf according to sediment type and structure. Side-scan sonar and our calculations indicate that the area affected by walrus feeding is in the order of thousands of square kilometers per year. Annual to long-term walrus bioturbation results in significant, large-scale changes in sediment and biological-community structure, and magnifies nutrient flux from sediment pore water to the water column by about two orders of magnitude over wide areas. The combined effects of walrus feeding must be placed in the context of long-term, regional climate changes and responses. Should sea ice continue to move northward as a result of climate change, the walrus' ecological role could be diminished or lost, the benthic ecosystem could be fundamentally altered and native subsistence hunters would be deprived of important resources.     

Diversity versus disparity and the radiation of modern cetaceans

Modern whales are frequently described as an adaptive radiation spurred by either the evolution of various key innovations (such as baleen or echolocation) or ecological opportunity following the demise of archaic whales. Recent analyses of diversification rate shifts on molecular phylogenies raise doubts about this interpretation since they find no evidence of increased speciation rates during the early evolution of modern taxa. However, one of the central predictions of ecological adaptive radiation is rapid phenotypic diversification, and the tempo of phenotypic evolution has yet to be quantified in cetaceans. Using a time-calibrated molecular phylogeny of extant cetaceans and a morphological dataset on size, we find evidence that cetacean lineages partitioned size niches early in the evolutionary history of neocetes and that changes in cetacean size are consistent with shifts in dietary strategy. We conclude that the signature of adaptive radiations may be retained within morphological traits even after equilibrium diversity has been reached and high extinction or fluctuations in net diversification have erased any signature of an early burst of diversification in the structure of the phylogeny.     

Coexisting living stromatolites and infaunal metazoans

Microbialites, bioaccretionary structures formed during the growth and metabolism of microorganisms (principally cyanobacteria) were the dominant lifeform in shallow late-Archean and Proterozoic oceans. During the Cambrian radiation of metazoan life, which began ~540 Mya, microbialite abundance and diversity further declined following a peak in the Mesoproterozoic. Notwithstanding contention, grazing and bioturbation effects of metazoans have been hypothesized as the dominant driver of modern microbialite scarcity. However, this metazoan–microbialite exclusion has not been fully explored in the few extant microbialites. Here we provide further evidence showing that living marine layered microbialites (stromatolites) coexist with a persistent assemblage of benthic macro-invertebrates, as has previously been demonstrated in some thrombolitic (clotted) microbialites. Surprisingly, these metazoans have active habits, such as burrowing, which should be expected to disrupt the layered matrix. As other studies have shown, through a network of burrows, metazoans can exploit local diurnal oxygen refugia within microbialites as well as escape predation. Our results, therefore, add novel evidence in support of the hypotheses that geologically, metazoans are not always incompatible with stromatolites, while ecologically, microbialites may act as micro-refugia for modern metazoans and historically have performed a similar inferred role in past ecosystems.     

Plant growth forms: an ecological and evolutionary perspective

Trees, shrubs, lianas and herbs have widely different mechanical architectures, which can also vary phenotypically with the environment. This review investigates how environmental effects, particularly mechanical perturbation, can influence biomechanical development in self-supporting and climbing growth forms. The bifacial vascular cambium is discussed in terms of its significance to growth form variation, ecology and evolution among extant plants, and during its appearance and early evolution. A key aspect of this developmental innovation concerned its potential for architectural and mechanical variation in response to environmental effects as well as optimizing hydraulic supply before the appearance of laminate leaves. Growth form diversity and its importance to past and present ecosystems are discussed in relation to both evolutionary constraints and ecological factors such as climatic change and atmospheric CO2 concentrations. We discuss how widely ranging growth forms such as climbers show a large range of developmental and phenotypic variation that has much to offer in understanding how the environment can modify plant development, particularly in terms of the bifacial vascular cambium. The broad approach we propose would benefit a wide range of studies from research into wood development to long-term ecological censuses of today's potentially changing ecosystems.     

Notes on an interspecific interaction between distant taxa: Observation of active cohabitation and communal nesting resources between Gehyra mutilata (Wiegmann, 1834) and two vespid wasp species

Tropical ecosystems are biodiverse and ecologically complex, supporting nearly half of the global herpetofauna. Ecological interactions play a role in the maintenance of species diversity with a range of ecological interactions occurring within and among species in tropical ecosystems. Incidental observations of ecological interactions provide an important contribution to the understanding of the ecology and natural history of species. Here, we report an interspecific interaction between distant taxa, the Four-Clawed Gecko (Gehyra mutilata) utilizing an active communal nesting site and shared nesting resources with two vespid wasp species (Eumeninae and Polistinae), where previous observations documented species utilizing abandoned nesting resources. This is the first known record of three species actively cohabitating in a shared nesting resource. This observation, with one or all species positively benefiting from the ecological interaction, likely indicates a mutualistic or commensal ecological interaction. Incidental observations such as the present study are likely to reveal that interspecific ecological interactions including communal nesting are more widespread than previously thought and the co-utilization of existing nesting resources presents an advantage for taxa. Observations such as these provide the basis for understanding the complex network of ecological interactions occurring in tropical rainforests.     

淡水鱼类功能多样性及其研究方法

目前,群落功能多样性备受生态学界关注,被认为是能解决生态问题的一种重要途径。我国对于群落功能多样性主要集中在植物群落和微生物群落,而在鱼类群落方面的研究几乎是空白。我国鱼类资源正面临着严重威胁,包括水坝建设导致的鱼类通道受阻、水库形成造成鱼类产卵场功能消失、过度捕捞、水质恶化和富营养化加重、外来种入侵等因素,导致渔业资源急剧衰退,水生生态系统功能下降。以淡水鱼类群落为例,对鱼类功能多样性的数据获取及处理分析与评价、测定指标及计算方法与研究难点等进行综述,以期为鱼类资源保护提供新的理论依据和切入点。