A recipe for scavenging in vertebrates – the natural history of a behaviour

Human-induced environmental change has caused widespread loss of species that support important functions for ecosystems and society. For example, vertebrate scavengers contribute to the functional health of ecosystems and provide services to agricultural landscapes by removing carcasses and associated pests. Widespread extirpation of native Australian mammals since the arrival of Europeans in Australia has removed many scavenging species from landscapes, while scavenging mammals such as European red foxes (Vulpes vulpes) have been introduced. In much of Australia, squamate reptiles are the largest native terrestrial scavengers remaining, where large native mammals are extinct and conservation management is being undertaken to remove invasive mammals. The contribution of reptiles to scavenging functions is not well understood. In this study, we investigated the ecosystem functions provided by large reptiles as scavengers to better understand how populations can be managed to support ecosystem services. We investigated the ecosystem services provided by vertebrate scavengers in Australian coastal mallee ecosystems, focusing on the heath goanna (Varanus rosenbergi), the only extant native terrestrial scavenger in the region. We carried out exclosure experiments, isolating the scavenging activity of different taxonomic groups to quantify the contribution of different taxa to scavenging services, specifically the removal of rat carcasses, and its impact on the occurrence of agriculturally damaging blowflies. We compared areas with different native and invasive scavenger communities to investigate the impact of invasive species removal and native species abundance on scavenging services. Our results indicated that vertebrate scavenging significantly contributes to carcass removal and limitation of necrophagous fly breeding in carcasses and that levels of removal are higher in areas associated with high densities of heath goannas and low densities of invasive mammals. Therefore, augmentation of heath goanna populations represents a promising management strategy to restore and maximize scavenging ecosystem services.

AimGlobally, urbanization is one of the most widespread, intense and ecologically destructive forms of landscape transformation, and it is often concentrated in coastal areas. Theoretically, species losses attributable to urbanization are predicted not to alter overall ecosystem function if functional redundancy (i.e. replacement of function by alternative species) compensates for such losses. Here, we test this expectation by measuring how coastal urbanization affects scavenger guilds on sandy beaches and whether changes in guild composition result either in an overall loss of scavenging efficiency, or in functional compensation under alternative guild structures, maintaining net ecosystem functioning.LocationFourteen beaches along the east coast of Australia with variable levels of urbanization.MethodsScavenging communities and rates of carrion removal were determined using motion‐triggered cameras at the beach‐dune interface.ResultsA substantial shift in the community structure of vertebrate scavengers was associated with gradients in urbanization. Iconic and functionally important raptors declined precipitously in abundance on urban beaches. Importantly, other vertebrates usually associated with urban settings (e.g. dogs, foxes, corvids) did not functionally replace raptors. In areas where < 15% of the abutting land had been developed into urban areas, carcass removal by scavengers was often complete, but always > 70%. Conversely, on beaches bordering coastal cities with < 40% of natural vegetation remaining, two‐thirds of fish carcasses remained uneaten by scavengers. Raptors removed 70–100% of all deployed fish carcasses from beaches with < 8% urban land cover, but this number dropped significantly with greater levels of urbanization and was not compensated by other scavenger species in urban settings.Main conclusionsThere is limited functional redundancy in vertebrate scavenger communities of sandy beach ecosystems, which impacts the system's capacity to mitigate the ecological consequences of detrimental landscape transformations.

The organization of ecological assemblages has important implications for ecosystem functioning, but little is known about how scavenger communities organize at the global scale. Here, we test four hypotheses on the factors affecting the network structure of terrestrial vertebrate scavenger assemblages and its implications on ecosystem functioning. We expect scavenger assemblages to be more nested (i.e. structured): 1) in species‐rich and productive regions, as nestedness has been linked to high competition for carrion resources, and 2) regions with low human impact, because the most efficient carrion consumers that promote nestedness are large vertebrate scavengers, which are especially sensitive to human persecution. 3) We also expect climatic conditions to affect assemblage structure, because some scavenger assemblages have been shown to be more nested in colder months. Finally, 4) we expect more organized assemblages to be more efficient in the consumption of the resource. We first analyzed the relationship between the nestedness of the scavenger assemblages and climatic variables (i.e. temperature, precipitation, temperature variability and precipitation variability), ecosystem productivity and biomass (i.e. NDVI) and degree of human impact (i.e. human footprint) using 53 study sites in 22 countries across five continents. Then, we related structure (i.e. nestedness) with its function (i.e. carrion consumption rate). We found a more nested structure for scavenger assemblages in regions with higher NDVI values and lower human footprint. Moreover, more organized assemblages were more efficient in the consumption of carrion. However, our results did not support the prediction that the structure of the scavenger assemblages is directly related to climate. Our findings suggest that the nested structure of vertebrate scavenger assemblages affects its functionality and is driven by anthropogenic disturbance and ecosystem productivity worldwide. Disarray of scavenger assemblage structure by anthropogenic disturbance may lead to decreases in functionality of the terrestrial ecosystems via loss of key species and trophic facilitation processes.

—The Snyder quarry is an Upper Triassic bonebed located in north-central New Mexico. The locality is stratigraphically high in the Petrified Forest Formation of the Chinle Group, and tetrapod biostratigraphy places it in the Revueltian land-vertebrate faunachron (mid-Norian in age: 210-215 Ma). This site has yielded the remains of a wide variety of organisms, ranging from terrestrial and aquatic vertebrates to aquatic invertebrates, as well as a substantial volume of charcoalified wood. A taphonomic analysis of both the biological material and the associated sediments indicates that the bonebed is the result of a catastrophic mass mortality event. The sediments of the bonebed contain rip-up clasts from the surrounding floodplain, a significant portion of the bone and wood is aligned, there is a high density of bones over a large area, and a moderate degree of hydraulic sorting of the skeletal material, indicating brief transport and rapid deposition of the bonebed. There is no evidence of abrasion on the bones, indicating that transport was minimal. The skeletal material is associated, with no evidence of substantial weathering or vertebrate scavenging, reflecting a rapid burial of partially dissociated carcasses. A survivorship curve constructed for the phytosaurs, the dominant taxonomic group, shows a thanatocoenosis that matches the biocoenosis for a reptile population. Scanning electron microscopy of the charcoalified wood reveals that the internal structure of the cell walls has been homogenized, and the reflectance of the charcoal is significantly higher than that of other forms of plant fossil preservation. These two pieces of data are evidence that the wood was burned in a moderate temperature (300-450°C) ground fire. Thus, both the sedimentological and biological data from the Snyder quarry best fit the scenario of a catastrophic, Late Triassic wildfire.

Carrion is a frequent but overlooked source of nutrients to the soil. The decomposition of carrion is accelerated by invertebrate scavengers, but the impact of the scavengers on below‐ground biota and its functions is scarcely known. We conducted a laboratory experiment to investigate the effects of the burying beetle Nicrophorus vespilloides on the soil community of a temperate broadleaved forest. We assembled microcosms from soil collected from an oak woodland and treated them with mouse Mus musculus carcasses and mating pairs of burying beetles (♀+♂) in a factorial design with control soils. We sampled independent replicates over time to investigate how the beetles affect soil microarthropods and microbial biomass (bacteria and fungi) in relation to soil pH and organic matter content. The beetle treatment initially reduced the total microbial biomass and abundance of major groups of microarthropods relative to the control soil. At the same time, organic matter increased in the beetle treatment and then dropped to the pre‐beetle level (i.e. soil baseline) at the end of the beetle breeding cycle (2 weeks). The rapid temporal changes in organic matter were mimicked by the relative abundances of the dominant microarthropod groups, with Oribatida relatively more abundant than Collembola and predaceous mites in the beetle treatment. The overall final effect of the beetle (relative to the laboratory control) on microarthropods was negative but the beetle kept these variables within the levels observed for freshly collected soil (baseline), while the final effect on pH was positive, and most likely driven by the surplus of nutrients from the carcass and biochemical changes triggered by the decomposition process. In nature, scavenging invertebrates are widespread. Our study demonstrates that beetles breeding in carcasses regulate the dynamics of key components of the soil food web, including microbial biomass, changes in the relative abundances of dominant microarthropods and soil organic matter and pH. Given the abundance of these beetles in nature, the study implies that the distribution of these beetles is a key driver of variation in soil nutrient cycling in woodlands. A free Plain Language Summary can be found within the Supporting Information of this article.

Recent analyses have suggested that extinction and origination rates exhibit long-range correlations, implying that the fossil record may be controlled by self-organized criticality or other scale-free internal dynamics of the biosphere. Here we directly test for correlations in the fossil record by calculating the autocorrelation of extinction [corrected] and origination rates through time. Our results show that extinction rates are uncorrelated beyond the average duration of a stratigraphic interval. Thus, they lack the long-range correlations predicted by the self-organized criticality hypothesis. In contrast, origination rates show strong autocorrelations due to long-term trends. After detrending, origination rates generally show weak positive correlations at lags of 5-10 million years (Myr) and weak negative correlations at lags of 10-30 Myr, consistent with aperiodic oscillations around their long-term trends. We hypothesize that origination rates are more correlated than extinction rates because originations of new taxa create new ecological niches and new evolutionary pathways for reaching them, thus creating conditions that favour further diversification.

Contribution of manipulable and non-manipulable environmental factors to trapping efficiency of invasive sea lamprey

Invertebrate scavengers of terrestrial and aquatic carrion include a broad and enormously diverse grouping of various arthropods, nematodes, and molluscs. By far the most functionally important in terrestrial ecosystems are insects, especially the flies, with crustaceans performing this role in marine ecosystems, although a multi-trophic foodweb of numerous invertebrate taxa can be found at most carcasses in most environments. The occurrence of different taxa, and the dynamics of their colonization and various competitive interactions varies across terrestrial and aquatic realms, and is greatly influenced by geographic region, climate, habitat, season, and other biotic factors. This sub-chapter focuses on the invertebrate use of carrion in both terrestrial and aquatic ecosystems, and highlights the role of different groups of invertebrates, and the biotic and abiotic factors that influence their occurrence and succession at decaying carcasses.

Comments on: Periodicity in the extinction rate and possible astronomical causes – comment on mass extinctions over the last 500 myr: an astronomical cause? (Erlykin <i>et al</i>.)

Two rival models of evolution in multispecies systems are tested against empirical species-level data. The two models are the Red Queen model of Van Valen as reformulated by Stenseth and Maynard Smith, which assumes that evolution is driven principally by biotic interactions, and the Stationary model of Stenseth and Maynard Smith, which assumes that evolution is propelled primarily by abiotic factors and will cease in the absence of changes in abiotic parameters. Testing refers to the models' predictions regarding the behavior of extinction and origination rates, and assumptions regarding equilibrium diversity and a constant effective environment. The data set includes the dates of origination and extinction for all coccolith, planktic foraminifer, and radiolarian species recorded in the Oligocene through Holocene, and all planktic diatom and silicoflagellate and ebridian species recorded in the Middle Miocene through Holocene in 111 DSDP sites of the low- to mid-latitude Pacific Ocean.The condition of stable specific age distribution over geologic time is met, which allows one to perform survivorship analysis on extinction rates. The best fit survivorship curve is a decreasing function of age for both coccolith and foraminifer species, and an increasing function of age for radiolarian species. Neither model predicts age dependence of the probability of extinction. The small disparity between these curves and age-independent curves for each group indicates, however, that an age-independent interpretation of extinction probability is a reasonable first approximation. Rates of origination are analyzed in terms of species accretion, introduced to represent the cumulative origination of species within a higher taxon as a function of the age or duration of the community. Accretion analysis indicates that the probability of accretion is both diversity-dependent and absolute time-dependent.The assumption of a constant effective environment is tested by polycohort analysis and nonparametric logistic regression analysis of true species cohorts. Both techniques indicate considerable variation in extinction probability over geologic time. When the predictions of the two evolutionary models are adjusted to take this variation into account, the results of both survivorship and accretion analysis seem to conform more closely to the predictions of the Red Queen than to the Stationary model. However, as the speed with which the effective environment changes is increased relative to speciation-extinction rates, it becomes increasingly difficult to differentiate patterns predicted by the two models. The assumption of equilibrium diversity can be neither corroborated nor rejected, since the species-level data are compatible with both an equilibrium and a nonequilibrium view of diversity behavior. Reservations concerning the basic assumptions of both models indicate an ultimate test requires that both models be reformulated to make precise and distinctive predictions under a varying effective environment.

We still have much to learn about the evolution of taxonomic diversity gradients through geologic time. For example, have latitudinal gradients always been as steep as they are now, or is this a phenomenon linked to some form of Cenozoic global climatic differentiation? The fossil record offers potential to address this sort of problem, and this study reconstructs latitudinal diversity gradients for the last (Tithonian) stage of the Jurassic period using marine bivalves. At this time of relative global warmth, bivalves were cosmopolitan in their distribution and the commonest element within macrobenthic assemblages.Analysis of 31 regional bivalve faunas demonstrates that Tithonian latitudinal gradients were present in both hemispheres, though on a much smaller magnitude than today. The record of the Northern Hemisphere gradient is more complete and shows a steep fall in values at the tropical/temperate boundary; the Southern Hemisphere gradient exhibits a more regular decline in diversity with increasing latitude.Tithonian latitudinal gradients were underpinned by a tropical bivalve fauna that comprises almost equal numbers of epifaunal and infaunal taxa. The epifaunal component was dominated by three pteriomorph families, the Pectinidae, Limidae and Ostreidae, that may be regarded as a long-term component of tropical bivalve diversity. Of the mixture of older and newer “heteroconch” families that formed the bulk of the infaunal component, the latter radiated spectacularly through the Late Cretaceous and Cenozoic to dominate tropical bivalve faunas at the present day. This pulse of heteroconch diversification, which was a major cause of the steepening of the bivalve latitudinal gradient, provides important evidence that rates of speciation may be negatively correlated with latitude.Nevertheless, we cannot exclude the possibility that elevated extinction rates in the highest latitudes also contributed to the marked steepening of bivalve latitudinal gradients over the last 150 Myr. Rates of extinction within epifaunal bivalve taxa appear to have been higher in these regions through the Cretaceous period, but this was largely before any significant global climatic deterioration. Infaunal bivalve clades have had differential success over this time period in the polar regions. Whereas, in comparison with the Tropics, heteroconchs are very much reduced in numbers today, the anomalodesmatans are much better represented, and the protobranchs have positively thrived. We are beginning to appreciate that low temperature per se may not be a primary cause of elevated rates of extinction. Food supply may be an equally important control on both rates of speciation and extinction; those bivalves that have been able to adapt to the extreme seasonality of food supply have flourished in the polar regions.

Correction for ‘Correlations in fossil extinction and origination rates through geological time’ by J. W. Kirchner and A. Weil (Proc. R. Soc. Lond. B 267 , 1301–1309. (doi: 10.1098/rspb.2000.1142)). The third sentence of the abstract is incorrect and should have appeared as: Here we directly test for correlations in the fossil record by calculating the autocorrelation of extinction and origination rates through time.

Recent Fourier analyses of fossil extinction data have indicated that the power spectrum of extinction during the Phanerozoic may take the form of 1/f noise, a result which, it has been suggested, could be indicative of the presence of `critical dynamics' in the processes giving rise to extinction. In this paper we examine extinction power spectra in some detail, using family-level data from two widely available compilations. We find that although the average form of the power spectrum roughly obeys the 1/f law, the spectrum can be represented more accurately by dividing it into two regimes: a low-frequency one which is well fit by an exponential, and a high-frequency one in which it follows a power law with a 1/f2 form. We give explanations for the occurrence of each of these behaviours and for the position of the crossover between them.

Many important phenomena in the plankton are driven by encounters among individuals. These encounters are mediated by the relative motion of zooplankters, either through the swimming ability of organisms, the small-scale hydrodynamic turbulence, or both. Through selected case studies, in this chapter, we illustrate how encounter rates influence the predator-prey interactions and reproduction, two of the major processes regulating the zooplankton population dynamics. Estimations on the encounter rates among zooplankters were made on the basis of the Gerritsen-Strickler and Rothschild-Osborn models, which consider non-turbulent and turbulent conditions, respectively. In a first case, we show how the predatory impact of siphonophores is over the fish larvae, in the southern Gulf of Mexico. In the absence of water turbulence, a predator encounters 38–40 prey in a day at surface waters, but under the influence of the wind, encounters can increase between 1.2 and 3.3 times depending on the wind velocity and prey speed. In a second case, we examined the encounters between a copepod predator and a cladoceran prey, the dominant groups in the meromictic lagoon of Clipperton atoll. Here, a predator can encounter a high number of prey (until 441) in a day, due to the high density of prey. Turbulence conditions enhance encounter rates, but even if encounters are high, it does not mean that a predator can ingest a high number of prey. In a third case, we analyzed the mate encounters of the holoplanktonic mollusk Firoloida desmarestia from the southern Gulf of Mexico, throughout an annual cycle. Results indicated that May is the high reproductive season, a period where a female can encounter 17 males in a day, under turbulent conditions. As F. desmarestia is a low abundant species, the role of wind-induced turbulence proved to be highly important in increasing encounters between mates. These case studies illustrate the importance of encounters among zooplankters in the growth and maintenance of populations in the plankton. Future field and experimental studies are needed to achieve a better understanding of the pelagic ecosystem dynamics.

Four Commentaries on the Pope’s Message on Climate Change and Income Inequality. IV. Pope Francis’ Encyclical Letter Laudato Si’, Global Environmental Risks, and the Future of Humanity.