Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains

Abstract

The golden eagle (Aquila chrysaetos) is one of the most important birds of prey in the Northern Hemisphere. This raptor is used to building large nests in high cliffs to which they return for several breeding years accumulating important amounts of their prey skeletal remains. This makes the golden eagle one of the major predators able to accumulate faunal remains in archaeological sites. Despite this fact, the taphonomic signature of golden eagles has not been properly characterized. Here we present the analysis of ingested and non-ingested faunal remains predated and accumulated by this raptor in two different nesting areas from the Iberian Peninsula. Results show how the faunal taxonomic record may vary depending on the ecological zone. Leporids and terrestrial carnivores are the best represented. The observed anatomical representation, breakage and bone surface modification patterns are discussed for different taxa. The taphonomic pattern varies depending on the type of prey and the origin of skeletal materials (non-ingested vs. pellets). Finally, after comparing our results with marks left by other predators, several characteristic features are noted to recognise golden eagles as agents of animal bones accumulations in the fossil record.     

Emerging aquatic insects as predators in terrestrial systems across a gradient of stream temperature in North and South America

1. Empirical and theoretical research over the past decade has demonstrated the widespread importance of aquatic subsidies to terrestrial food webs. In particular, adult aquatic insects that emerge from streams and lakes are prey for terrestrial predators. While variation in the magnitude of this subsidy is clearly important, the potential top‐down effects of the predatory adults of some aquatic insects in terrestrial food webs are largely unknown. 2. I used published data on benthic insect density (as a proxy for emergence) in North and South America to explore how the proportion of benthic insects that are predatory as adults varies across a gradient of mean annual stream temperature. 3. The proportion of benthic insects that are predatory as adults varied widely across sites (0–12% by abundance; 0–86% by biomass). There was a positive relationship between mean annual stream temperature and the proportion of predatory adults across all sites, driven largely by the greater abundance/biomass of predatory taxa (e.g. odonates), relative to non‐predators (e.g. midges, mayflies, caddisflies), in tropical than in temperate streams. 4. The ‘trophic structure’ (i.e. the proportion of predators) of emerging adult aquatic insects is an understudied source of variation in aquatic–terrestrial interactions. Incorporation of trophic structure in future studies is needed to understand how future modification of fresh waters may affect adjacent terrestrial food webs through both bottom‐up and top‐down effects.     

Herbivore damage increases avian and ant predation of caterpillars on trees along a complete elevational forest gradient in Papua New Guinea

Signals given off by plants to alert predators to herbivore attack may provide exciting examples of coevolution among organisms from multiple trophic levels. We examined whether signals from mechanically damaged trees (simulating damage by herbivores) attract predators of insects along a complete elevational rainforest gradient in tropical region, where various predators are expected to occur at particular elevational belts. We studied predation of artificial caterpillars on trees with and without ‘herbivorous’ damage; as well as diversity and abundances of potential predators at eight study sites along the elevational gradient (200–3700 m a.s.l.). We focused on attacks by ants and birds, as the main predators of herbivorous insect. The predation rate decreased with elevation from 10% d^?1 at 200 m a.s.l. to 1.8% d^?1 at 3700 m a.s.l. Ants were relatively more important predators in the lowlands, while birds became dominant predators above 1700 m a.s.l. Caterpillars exposed on trees with herbivorous damage were attacked significantly more than caterpillars exposed on trees without damage. Results suggest that relative importance of predators varies along elevational gradient, and that observed predation rates correspond with abundances of predators. Results further show that herbivorous damage attracts both ants and birds, but its effect is stronger for ants.     

Phylogenetic analysis of vertebrate kininogen genes

Kininogens, the precursors of bradykinins, vary extremely in both structure and function among different taxa of animals, in particular between mammals and amphibians. This includes even the most conserved bradykinin domain in terms of biosynthesis mode and structure. To elucidate the evolutionary dynamics of kininogen genes, we have identified 19 novel amino acid sequences from EST and genomic databases (for mammals, birds, and fishes) and explored their phylogenetic relationships using combined amino acid sequence and gene structure as markers. Our results show that there were initially two paralogous kininogen genes in vertebrates. During their evolution, the original gene was saved with frequent multiplication in amphibians, but lost in fishes, birds, and mammals, while the novel gene was saved with multiple functions in fishes, birds, and mammals, but became a pseudogene in amphibians. We also propose that the defense mechanism against specific predators in amphibian skin secretions has been bradykinin receptor dependent. Our findings may provide a foundation for identification and structural, functional, and evolutionary analyses of more kininogen genes and other gene families.     

Functional Invertebrate Prey Groups Reflect Dietary Responses to Phenology and Farming Activity and Pest Control Services in Three Sympatric Species of Aerially Foraging Insectivorous Birds

Farming activity severely impacts the invertebrate food resources of farmland birds, with direct mortality to populations of above-ground arthropods thorough mechanical damage during crop harvests. In this study we assessed the effects of phenological periods, including the timing of harvest, on the composition and biomass of prey consumed by three species of aerial insectivorous birds. Common Swifts Apus apus, Barn Swallows Hirundo rustica and House Martins Delichon urbica breed sympatrically and most of their diet is obtained from agricultural sources of invertebrate prey, especially from oil-seed rape crops. We categorized invertebrate prey into six functional groups, including oil-seed rape pests; pests of other arable crops; other crop-provisioned taxa; coprophilous taxa; and taxa living in non-crop and mixed crop/non-crop habitats. Seasonality impacted functional groups differently, but the general direction of change (increase/decrease) of all groups was consistent as indexed by prey composition of the three aerial insectivores studied here. After the oil-seed rape crop harvest (mid July), all three species exhibited a dietary shift from oil-seed rape insect pests to other aerial invertebrate prey groups. However, Common Switfts also consumed a relative large quantity of oil-seed rape insect pests in the late summer (August), suggesting that they could reduce pest insect emigration beyond the host plant/crop. Since these aerially foraging insectivorous birds operate in specific conditions and feed on specific pest resources unavailable to foliage/ground foraging avian predators, our results suggest that in some crops like oil-seed rape cultivations, the potential integration of the insectivory of aerial foraging birds into pest management schemes might provide economic benefits. We advise further research into the origin of airborne insects and the role of aerial insectivores as agents of the biological control of crop insect pests, especially the determination of depredation rates and the cascading effects of insectivory on crop damage and yield.     

Arboreal tropical forest vertebrates: current knowledge and research trends

We review the ecology and specialized methods required for studying arboreal mammals, birds, reptiles and amphibians, and use faunal checklists from 12 tropical wet forest sites and an analysis of all articles published during the past ten years in 14 major journals to assess current knowledge and general research trends for these groups. The percentage of arboreal vertebrates was remarkably similar at the different sites (76.2 ± 3.9%). Birds were the most arboreal group and amphibians and reptiles the least. The review of journals showed that primates were overwhelmingly the most studied group (336 papers), followed by bats (105), passeriform birds (73) and rodents (55). Judging by their portion of the arboreal vertebrate community and the number of papers surveyed, birds and amphibians and reptiles are vastly understudied compared to mammals, but this is largely due to the great number of primate studies. The number of publications on arboreal vertebrates has remained relatively stable over the last 10 years for all taxa except primates, which have seen a growth in publications. Canopy vertebrates from Brazil had by far the most publications (120), followed by Madagascar (61), Costa Rica (55) and Indonesia (42). We conclude by highlighting the priorities we see for future studies on tropical canopy vertebrates.     

Nest-site competition between bumblebees (Bombidae), social wasps (Vespidae) and cavity-nesting birds in Britain and the Western Palearctic

Capsule There is no evidence of widespread significant nest-site competition in Britain or the Western Palearctic between cavity-nesting birds and bumblebees or social wasps.

Aims To investigate competition between cavity-nesting birds and bumblebees and wasps, particularly the range-expanding Tree Bumblebee, Saxon Wasp and European Hornet in Britain, and review evidence throughout the Western Palearctic.

Methods We compared field data from English and Polish studies of tits and woodpeckers breeding in nest-boxes and/or tree holes to assess nest-site competition with bumblebees and wasps. We reviewed the literature quantifying nest-site competition between birds and these insects in the Western Palearctic.

Results Bumblebees and wasps are capable of usurping small passerines from nests. In England, these insects commandeered a mean annual 4.1% of tit nests initiated in nest-boxes; occurrence of hornets showed a long-term increase, but not other wasps or bumblebees. Across the Western Palearctic, insect occupation of nest-boxes was generally low, and was lower in England than in Poland. No insects were discovered in tree cavities, including those created by woodpeckers (Picidae).

Conclusion Nest-site competition between cavity-nesting birds and bumblebees and wasps appears to be a ‘nest-box phenomenon’, which may occasionally interfere with nest-box studies, but appears negligible in natural nest-sites.     

The diet of feral cats on islands: a review and a call for more studies

Cats are among the most successful and damaging invaders on islands and a significant driver of extinction and endangerment. Better understanding of their ecology can improve effective management actions such as eradication. We reviewed 72 studies of insular feral cat diet from 40 islands worldwide. Cats fed on a wide range of species from large birds and medium sized mammals to small insects with at least 248 species consumed (27 mammals, 113 birds, 34 reptiles, 3 amphibians, 2 fish and 69 invertebrates). Three mammals, 29 birds and 3 reptiles recorded in the diet of cats are listed as threatened by the IUCN. However, a few species of introduced mammals were the most frequent prey, and on almost all islands mammals and birds contributed most of the daily food intake. Latitude was positively correlated with the predation of rabbits and negatively with the predation of reptiles and invertebrates. Distance from landmass was positively correlated with predation on birds and negatively correlated with the predation of reptiles. The broad range of taxa consumed by feral cats on islands suggests that they have the potential to impact almost any native species, even the smallest ones under several grams, that lack behavioral, morphological or life history adaptations to mammalian predators. Insular feral cat??s reliance on introduced mammals, which evolved with cat predation, suggests that on many islands, populations of native species have already been reduced.     

Birds have no impact on folivorous insect guilds on a montane willow

We examined the impact of avian predators on the community of insects feeding on Lemmon's willow, Salix lemmonii , at Carman Valley, California. Salix lemmonii supports a diverse assemblage of insect species that feed in either a concealed or an exposed manner on leaves by mining, tying, curling, galling, skeletonizing, or chewing. We tested the hypothesis that there are guild specific differences in the effects of avian predators on concealed and exposed-feeding folivores. We compared the feeding damage and frequency of each guild in the presence and absence of avian predators. Contrary to most studies of the effects of avian predators on folivorous insects, birds failed to reduce significantly the frequency of or feeding damage caused by exposed-feeding insects during 1999 and 2000. Chewing insects, which accounted for the majority of foliar damage by exposed-feeding insects, inflicted slightly more damage and skeletonizing insects slightly less damage in the presence of birds. We also observed no reduction in the frequency of concealed-feeding insects in the presence of avian predators; rather three out of four guilds were slightly more common in the presence of avian predators. While the abundance of foliage gleaning birds at the study site was slightly lower than the average for the past nine years, we interpret our results to indicate a true lack of an effect of avian predators on the community of folivorous insects on S. lemmonii. Future inquiry into the effects of avian predators on folivorous insects should focus on determining under what circumstances birds will or will not suppress guilds of folivorous insects.     

Taphonomic study of leporid remains accumulated by the Spanish Imperial Eagle (Aquila adalberti)

Distinguishing leporid bones accumulated by different agents such as diurnal raptors, owls, mammals and humans is essential to gain an understanding of not only human subsistence activities but also past ecology. This is particularly relevant in Iberian Palaeolithic sites where leporid remains usually constitute the most abundant taxon. Among diurnal raptors the Spanish Imperial eagle (Aquila adalberti) has been one of the most important leporid predators throughout the Iberian Peninsula. In order to investigate the taphonomic signature of this raptor, rabbit remains from 79 pellets were examined. Results show a high proportion of distal elements of the limb bones and skull. Compared with other diurnal birds of prey, the assemblages produced by this species appear to show a higher degree of breakage and corrosion from digestion. These results place this predator within a category similar to the small mammal carnivores (category 5) in terms of skeletal element abundance, breakage and digestion. It is hoped that these data will enable analysts to identify leporid fossils accumulated by the Spanish Imperial Eagle in archaeological assemblages.     

Birds and prehistoric agriculture: The New Mexican pueblos

Analysis of faunal remains from prehistoric southwestern pueblo sites reveals certain commonly occurring species of mammals and birds, the latter including ducks and geese, hawks and eagles, American Kestrel, Mourning Dove, owls, Horned Lark, Common Raven, and jays. This phenomenon is examined with the analysis of bird bones from four New Mexican pueblos located in north-central and west-central New Mexico. Recovered faunal remains date to specific periods between A.D. 1250 and the present. Over 70 species of birds represent various ecological situations; many of these species presently do not occur near the pueblos and a few occur only in the extreme southeastern portion of the state. Marsh and riparian areas along the river systems and richer grasslands on the floodplains would account for most of these species occurring near the sites prehistorically. However, rather than attribute unusual species to climatic change, trade, or long-distance hunting expeditions by the Indians, an alternative explanation is offered which attributes high species diversity to areas of human disturbance. Disturbed areas such as agricultural fields can increase species diversity in an area and aid in the range extension of many animals. The primary factor in this situation, besides the crops themselves, is believed to be insects, which attract other animals, particularly birds and their predators. In addition, field irrigation systems act to extend riparian growth from the river across the floodplain, creating habitats for these animals. Finally, certain groups of birds and mammals have developed a natural attraction to human settlements and agricultural areas and these include species whose remains commonly occur in prehistoric pueblo sites. The avifauna from these sites also reveals information on the prehistoric use of birds as inferred from bone modification, cut marks, burials, and articulated wings.     

An updated review of the middle Eocene avifauna from the Geiseltal (Germany), with comments on the unusual taphonomy of some bird remains

The brown coal mines of the Geiseltal in Germany were among the most important middle Eocene fossil localities and constitute the reference sites for the Geiseltalian stage of the European Land Mammal Mega Zones. Here, an updated review of the Geiseltal avifauna is given. Thirteen species are represented by diagnostic bones and can be referred to avian higher-level taxa, but various indeterminable albeit distinct fossils indicate a higher diversity of the Geiseltal avifauna. The majority of the Geiseltal fossils belong to taxa, which are well known from early to mid-Ypresian fossil sites of Europe and North America. Some of the species from the Geiseltal are larger than their early Eocene relatives, but it remains elusive whether this indicates an evolutionary size increase in some avian lineages or reflects different palaeohabitats of the involved fossil sites. The Geiseltal avifauna exhibits some taphonomic peculiarities that have not yet been addressed. Most bird fossils consist of isolated bones or partial skeletons and here it is hypothesized that at least some of the fragmentary specimens represent feeding remains of crocodilians. For the first time, medullary bone, which is indicative of breeding females, is reported for birds from the Geiseltal. The comparative abundance of the coliiform species Eoglaucidium pallas is remarkable, and either this species was extremely abundant in the Geiseltal ecosystem or it had ecological preferences, which favored its preservation.     

The species diversity of New Zealand forest birds: Some possible consequences of the modification of beech forests

The literature on forest birds in New Zealand is examined to see if the consequences of forest modification can be predicted. Forty‐four samples of birds, from North, South, and Stewart Islands, are classified according to four habitats of origin: virgin forest, modified forest, exotic forest, and suburban bush and gardens. An index of bird species diversity (BSD) is calculated for comparison, and where possible densities are tabulated.

Limited samples from modified forests in Westland suggest that the number of species is relatively smaller but the bird density is higher after modification. There do not seem to be any published quantitative data about birds in Westland exotic forests.

Data from the whole of New Zealand show that bird density increases as forest habitats change from virgin, to modified, to exotic, to suburban bush and gardens. BSD and equitability do not differ among these habitats. The number of species of native birds and their diversity decline as the habitat is more extensively altered, but their total density tends to remain constant. More productive forest habitats have the same BSD as other habitats. Diversity seems to be limited by the small variety of bird species available, and the more productive habitats could probably support more species. All habitats had a much higher diversity of birds before colonisation by the European.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.     

The predators of insects

Abstract. 1. Insect–insectivore trophic relations were reviewed using presence–absence data from sixty-one invertebrate-dominated food webs and fifteen food webs from Briand's (1983) original forty web collection. From counts of prey links in higher taxa (orders, classes, phyla), six phyla and thirteen classes of non-insect insectivores and fourteen orders of insect predators and prey were found. 2. Detritus-based habitats (phytotelmata, felled logs, carcasses, dungpads) harboured fewer orders of insects, that interact with other insects, than webs from grazer-based (host plants, some galls) and mixed-based systems (aquatic webs). Consumer–resource networks of higher insect taxa in these webs shared several features found in some species-level biological networks: the trend was towards few pairs of strong asymmetrical links, several weak links and many null interactions. 3. From counts of insect predator–insect prey links, hymenopterans as terrestrial predators and parasitoids interacted with the most number of higher insect taxa. Hymenopterans were also linked as prey more often than other terrestrial insects. In freshwater habitats, plecopterans were linked as predators more often than other aquatic taxa, whereas dipterans were listed as prey more often than other insects. 4. Dipterans were linked in the diets of non-insect insectivores from seven of eight common taxonomic classes. Arachnids were identified as insect predators by food web researchers in the largest number of webs, followed by passerine birds and cyprinodont fishes. From analysis of prey links at the ordinal level, predaceous insects were less polyphagous than other predators (other ectotherms and endotherms). 5. Analysis of chain lengths, as expected, showed that insect prey occupied mostly lowermost trophic levels, non-insect insectivores were found mostly at uppermost trophic levels, and predaceous insects were found mostly at intermediate trophic levels across most habitats. 6. This analysis offers evidence that insects are not just occupying intermediate trophic levels in some communities. Indeed, some taxa feed at the upper ends of long food chains, for example eupelmids in galls, staphylinids in carcasses, and perlid plecopterans in streams.     

Flora and fauna assemblages vary with local topography in a subtropical eucalypt forest

Local‐scale spatial patterning (over hundreds of metres) in terrestrial assemblages was investigated by sampling a variety of organisms within a 400 ha eucalypt forest area in the lowlands of south‐east Queensland. Organisms were trees, shrubs, birds, insects extracted from the litter layer, and insects caught in pitfall traps. Each group was sampled using a standardized methodology, and the component taxa were counted and identified to a level commonly used in ecological studies of that organism – varying from species to order levels. Sites adjacent to drainage lines or ephemeral streams were biotically more similar to one another than they were to paired upslope sites 100–200 m distant and 15–35 m higher in altitude. This phenomenon occurred irrespective of the level of taxonomic resolution or type of organism. Within each taxonomic group, some components were mainly riparian, while others were more characteristic of upslope sites. Characteristically riparian taxa included trees in the genus Glochidion, the shrub genus Leptospermum, birds in the Pachycephalidae and Meliphagidae families, and litter invertebrates in the order Acarina. Upslope taxa included shrubs in the Rutaceae, birds in the Artamidae, and ants. Within the groups of trees, birds and litter invertebrates, more taxa were characteristic of riparian than upslope sites. Local scale biotic patterns were more strongly correlated with altitude than with measured soil characteristics; however, microtopographical differences would also be highly correlated with a large suite of covarying environmental features. The patterns of diversity and the implications for survey design and conservation are discussed.     

Complex impact of an invasive crayfish on freshwater food webs

Invasive alien species can have complex effects on native ecosystems, and interact with multiple components of food webs, making it difficult a comprehensive quantification of their direct and indirect effects. We evaluated the relationships between the invasive crayfish, Procambarus clarkii, amphibian larvae and predatory insects, to quantify crayfish impacts on multiple levels of food webs, and to evaluate whether crayfish predation of aquatic insects has indirect consequences for their preys. We used pipe sampling to assess the abundance of crayfish, amphibian larvae and their major predators (Ditiscidae, Notonectidae and larvae of Anisoptera) in invaded and uninvaded ponds within a human dominated landscape. We disentangled the multivariate effects of P. clarkii on different components of food web through a series of constrained redundancy analyses. The crayfish had a negative, direct impact on both amphibian communities and their predators. Amphibian abundance was negatively related to both predators. However, the negative, direct effects of crayfish on amphibians were much stronger than predation by native insects. Our results suggest that this crayfish impacts multiple levels of food webs, disrupting natural prey-predator relationships.     

Bone Fragments and Bone Grease Manufacturing: A Review of their Archaeological Use and Potential

Abstract

The presence of small bone fragments in the archaeological record has been frequently used to infer the activities of bone grease and/or soup manufacturing. This article explores the possibility that the activity or activity-complexes responsible for the bone fragments might be manipulated to account for additional archaeological data. Emphasis is placed on building predictive models against which the data from three Northern Plains-Upper Midwest archaeological sites will be compared. Results indicate that such an approach provides more information about the cultural activities being conducted at a site than does the other approach.     

Nestedness as a conservation tool: plants and birds of oak-hazel woodland in Sweden

Nestedness was examined for vascular plants and birds in the centres and edges of 26 sites of ancient oak-hazel woodland in Sweden. Both taxa exhibited significant nestedness in site centres and for whole sites, but not at the edges for birds. Woodland ranks of nestedness differed between plants and birds. Rank of nestedness of birds, but not of plants, depended on area. Horizontal habitat structure affected nestedness of both plants and birds. Mobility appears decisive for creating rank differences between sites for various taxa. High mobility may also explain a greater edge effect in birds from allochtonous, more or less transient individuals. Nestedness in relation to mobility, particularly at edges, should be of theoretical interest. The possible use of nestedness patterns in conservation makes further analyses urgent for less mobile taxa.     

Land‐use effects on terrestrial consumers through changed size structure of aquatic insects

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