The ones that come ready made:The identification and use of Sus tusks as tools at prehistoric cave sites in Malaysia

Evidence for osseous technologies has featured in excavation reports from Southeast Asia for almost a century and from archaeological deposits as old as 43,000 years BP. However, in contrast to the significance that is placed on this technology in other parts of the world, until recently, Southeast Asian assemblages have drawn only very limited attention. Concentrating on evidence from Malaysia, the current paper examines one element of this inventory of tools: the deliberate modification of pig canines and the means by which such alteration can be distinguished from patterns of natural tooth wear. Particular attention is paid to the bearded pig (Sus barbatus), as it is one of the two species of wild boar in Malaysia whose tusks are most likely to have been used by prehistoric toolmakers. Reference is also made to wider, regional ethnographic examples of known tusk implements and their accredited uses to further assist in the identification process. Distinguishing criteria for worked tusk are formulated according to the type and extent of modification. These criteria are then applied to archaeological specimens recovered from two prehistoric cave sites in Malaysia, Gua Bintong and Niah Cave.

The extended phenotype of Scots pine Pinus sylvestris structures the understorey assemblage

Heritable variation in plant secondary compounds in dominant species has been hypothesised to effect ecosystem function and the structure of associated assemblages of plants, microbes and animals. The functioning of this extended phenotype in relation to the understorey vegetation composition was tested within a boreal forest system dominated by Pinus sylvestris which contains a range of monoterpenes, the composition of which is largely under genetic control. A variance partitioning approach was adopted to identify the relative importance of tree chemistry, environment, spatial location and tree architecture in controlling the distribution of species in the ground flora under individual trees. The monoterpene composition of the pine needles appeared to contribute significantly to controlling understorey vegetation composition, but was less important than environmental factors, though similar to spatial factors. Thus there appears to be a link between variation in the chemical composition of the single, dominant tree species within this system and the pattern of occurrence and abundance in other species at the same trophic level.

Fractal species distributions do not produce power-law species-area relationships

We derive the species-area relationship (SAR) expected from an assemblage of fractally distributed species. If species have truly fractal spatial distributions with different fractal dimensions, we show that the expected SAR is not the classical power-law function, as suggested recently in the literature. This analytically derived SAR has a distinctive shape that is not commonly observed in nature: upward-accelerating richness with increasing area (when plotted on log-log axes). This suggests that, in reality, most species depart from true fractal spatial structure. We demonstrate the fitting of a fractal SAR using two plant assemblages (Alaskan trees and British grasses). We show that in both cases, when modelled as fractal patterns, the modelled SAR departs from the observed SAR in the same way, in accord with the theory developed here. The challenge is to identify how species depart from fractality, either individually or within assemblages, and more importantly to suggest reasons why species distributions are not self-similar and what, if anything, this can tell us about the spatial processes involved in their generation.

The geographical structure of British bird distributions:diversity, spatial turnover and scale

1. Using data on the spatial distribution of the British avifauna, we address three basic questions about the spatial structure of assemblages: (i) Is there a relationship between species richness (alpha diversity) and spatial turnover of species (beta diversity)? (ii) Do high richness locations have fewer species in common with neighbouring areas than low richness locations?, and (iii) Are any such relationships contingent on spatial scale (resolution or quadrat area), and do they reflect the operation of a particular kind of species-area relationship (SAR)?

2. For all measures of spatial turnover, we found a negative relationship with species richness. This held across all scales, with the exception of turnover measured as beta (sim).

3. Higher richness areas were found to have more species in common with neighbouring areas.

4. The logarithmic SAR fitted better than the power SAR overall, and fitted significantly better in areas with low richness and high turnover.

5. Spatial patterns of both turnover and richness vary with scale. The finest scale richness pattern (10 km) and the coarse scale richness pattern (90 km) are statistically unrelated. The same is true of the turnover patterns.

6. With coarsening scale, locations of the most species-rich quadrats move north. This observed sensitivity of richness 'hotspot' location to spatial scale has implications for conservation biology, e.g. the location of a reserve selected on the basis of maximum richness may change considerably with reserve size or scale of analysis.

7. Average turnover measured using indices declined with coarsening scale, but the average number of species gained or lost between neighbouring quadrats was essentially scale invariant at 10-13 species, despite mean richness rising from 80 to 146 species (across an 81-fold area increase). We show that this kind of scale invariance is consistent with the logarithmic SAR.

Bird diversity and environmental gradients in Britain:a test of the species-energy hypothesis

1. We tested the species diversity-energy hypothesis using the British bird fauna. This predicts that temperature patterns should match diversity patterns. We also tested the hypothesis that the mechanism operates directly through effects of temperature on thermoregulatory loads; this further predicts that seasonal changes in temperature cause matching changes in patterns of diversity, and that species' body mass is influential.

2. We defined four assemblages using migration status (residents or visitors) and season (summer or winter distribution). Records of species' presence/absence in a total of 2362, 10 x 10-km, quadrats covering most of Britain were used, together with a wide selection of habitat, topographic and seasonal climatic data.

3. We fitted a logistic regression model to each species' distribution using the environmental data. We then combined these individual species models mathematically to form a diversity model. Analysis of this composite model revealed that summer temperature was the factor most strongly associated with diversity.

4. Although the species-energy hypothesis was supported, the direct mechanism, predicting an important role for body mass and matching seasonal patterns of change between diversity and temperature, was not supported.

5. However, summer temperature is the best overall explanation for bird diversity patterns in Britain. It is a better predictor of winter diversity than winter temperature. Winter diversity is predicted more precisely from environmental factors than summer diversity.

6. Climate change is likely to influence the diversity of different areas to different extents; for resident species, low diversity areas may respond more strongly as climate change progresses. For winter visitors, higher diversity areas may respond more strongly, while summer visitors are approximately neutral.

Holocene “turn-on” and evolution of the Southern Great Barrier Reef: Revisiting reef cores from the Capricorn Bunker Group

Extensive drilling of the Great Barrier Reef (GBR) in the 70s and 80s illuminated the main factors controlling reef growth during the Holocene. However, questions remain about: (1) the precise nature and timing of reef "turnon" or initiation, (2) whether consistent spatio-temporal patterns occur in the bio-sedimentologic response of the reef to Holocene sea-level rise then stability, and (3) how these factors are expressed in the context of the different evolutionary states (juvenile-mature-senile reefs). Combining 21 new C14-AMS and 146 existing recalibrated radiocarbon and U/Th ages, we investigated the detailed spatial and temporal variations in sedimentary facies and coralgal assemblages in fifteen cores across four reefs (Wreck, Fairfax, One Tree and Fitzroy) from the Southern GBR. Our newly defined facies and assemblages record distinct chronostratigraphic patterns in the cores, displaying both lateral zonation across the different reefs and shallowing upwards sequences, characterised by a transition from deep (Porites/faviids) to shallow (Acropora/Isopora) coral types. The revised reef accretion curves show a significant lag period, ranging from 0.7-2 ka, between flooding of the antecedent Pleistocene substrate and Holocene reef turn-on. This lag period and dominance of more environmentally tolerant early colonizers (e.g., domal Porites and faviids), suggests initial conditions that were unfavourable for coral growth. We contend that higher input of fine siliciclastic material from regional terrigenous sources, exposure to hydrodynamic forces and colonisation in deeper waters are the main factors influencing initially reduced growth and development. All four reefs record a time lag and we argue that the size and shape of the antecedent platform is most important in determining the duration between flooding and recolonisation of the Holocene reef. Finally, our study of Capricorn Bunker Group Holocene reefs suggests that the size and shape of the antecedent substrate has a greater impact on reef evolution and final evolutionary state (mature vs. senile), than substrate depth alone. 

Late Quaternary climatic record from ODP Site 705 in the Northern Indian Ocean

The uppermost 500cm sedimentary core from ODP site located at the Eastern flank of Najareth bank in the Northern Indian Ocean has yielded altogether twenty four species of planktonic foraminifera. Among all these species, Globorotalia menardii has been found to be consistently dominant in the faunal assemblages from most of the samples. The 18O measured on the tests of Globorotalia menardii from all levels help in precisely working out the sediment accumulation rates at different isotopic stages, and deciphering the change in climate in the Late Quaternary as well.

John Birks: Pioneer in quantitative palaeoecology

We describe the career of John Birks as a pioneering scientist who has, over a career spanning five decades, transformed palaeoecology from a largely descriptive to a rigorous quantitative science relevant to contemporary questions in ecology and environmental change. We review his influence on students and colleagues not only at Cambridge and Bergen Universities, his places of primary employment, but also on individuals and research groups in Europe and North America. We also introduce the collection of papers that we have assembled in his honour. The papers are written by his former students and close colleagues and span many of the areas of palaeoecology to which John himself has made major contributions. These include the relationship between ecology and palaeoecology, late-glacial and Holocene palaeoecology, ecological succession, climate change and vegetation history, the role of palaeoecological techniques in reconstructing and understanding the impact of human activity on terrestrial and freshwater ecosystems and numerical analysis of multivariate palaeoecological data.

A spatio-temporal contrast of the predatory impact of an invasive freshwater crustacean

Aim: Impacts of invasive species may vary across invasion gradients, owing to trait-based sorting of individuals through dispersal: those aggregating at invasion fronts may be more aggressive and voracious. We examine, in the field and laboratory, variation in the predatory impacts of an invasive Ponto-Caspian crustacean Hemimysis anomala G.O. Sars, 1907 at two sites along a spatio-temporal gradient of invasion.

Location: Republic of Ireland.

Methods: We used reciprocal transplant field-deployed mesocosms to compare predation rates of invasion front and well-established H. anomala on natural zooplankton assemblages. In the laboratory, we measured the functional response (relationship between predation rate and prey supply) of H. anomala from both sites, for a per capita mechanistic comparison of predation efficiency. We also assessed prey selectivity of H. anomala in the mesocosm experiments to further compare feeding behaviour. Finally, we used a correlative approach to assess the community impact of H. anomala across sites, including a nearby uninvaded site, by comparing zooplankton diversities and densities.

Results: Invasion front H. anomala had higher predation rates than well-established H. anomala at high in situ zooplankton densities. Invasion front H. anomala also had higher functional responses - in particular showing higher 'attack rates' - indicating a heightened ability to locate and capture prey. Prey selectivity was consistent across the spatio-temporal contrast, with positive selection for cladocerans. Zooplankton diversity and density declined with time since H. anomala invasion, both being maximal at the uninvaded site.

Main conclusions: Our study, for the first time, (1) reveals differences in predatory per capita effects and associated behavioural traits between two sites along a spatio-temporal invasion gradient and (2) shows a negative community-level impact of the invasive H. anomala in natural water bodies. Further spatio-temporal comparisons of predatory per capita effects of invaders are needed to assess the generality of these results.

Deglacial mesophotic reef demise on the Great Barrier Reef

Submerged reefs are important recorders of palaeo-environments and sea-level change, and provide a substrate for modern mesophotic (deep-water, light-dependent) coral communities. Mesophotic reefs are rarely, if ever, described from the fossil record and nothing is known of their long-term record on Great Barrier Reef (GBR). Sedimentological and palaeo-ecological analyses coupled with 67 14C AMS and U–Th radiometric dates from dredged coral, algae and bryozoan specimens, recovered from depths of 45 to 130 m, reveal two distinct generations of fossil mesophotic coral community development on the submerged shelf edge reefs of the GBR. They occurred from 13 to 10 ka and 8 ka to present. We identified eleven sedimentary facies representing both autochthonous (in situ) and allochthonous (detrital) genesis, and their palaeo-environmental settings have been interpreted based on their sedimentological characteristics, biological assemblages, and the distribution of similar modern biota within the dredges. Facies on the shelf edge represent deep sedimentary environments, primarily forereef slope and open platform settings in palaeo-water depths of 45–95 m. Two coral–algal assemblages and one non-coral encruster assemblage were identified: 1) Massive and tabular corals including Porites, Montipora and faviids associated with Lithophylloids and minor Mastophoroids, 2) platy and encrusting corals including Porites, Montipora and Pachyseris associated with melobesioids and Sporolithon, and 3) Melobesiods and Sporolithon with acervulinids (foraminifera) and bryozoans. Based on their modern occurrence on the GBR and Coral Sea and modern specimens collected in dredges, these are interpreted as representing palaeo-water depths of < 60 m, < 80–100 m and > 100 m respectively. The first mesophotic generation developed at modern depths of 85–130 m from 13 to 10.2 ka and exhibit a deepening succession of < 60 to > 100 m palaeo-water depth through time. The second generation developed at depths of 45–70 m on the shelf edge from 7.8 ka to present and exhibit stable environmental conditions through time. The apparent hiatus that interrupted the mesophotic coral communities coincided with the timing of modern reef initiation on the GBR as well as a wide-spread flux of siliciclastic sediments from the shelf to the basin. For the first time we have observed the response of mesophotic reef communities to millennial scale environmental perturbations, within the context of global sea-level rise and environmental changes.

Predator-free space, functional responses and invasions.

1. Predator–prey interactions are mediated by the structural complexity of habitats, but disentangling the many facets of structure that contribute to this mediation remains elusive. In a world replete with altered landscapes and biological invasions, determining how structure mediates the interactions between predators and novel prey will contribute to our understanding of invasions and predator–prey dynamics in general.
2. Here, using simplified experimental arenas, we manipulate predator-free space, whilst holding surface area and volume constant, to quantify the effects on predator–prey interactions between two resident gammarid predators and an invasive prey, the Ponto-Caspian corophiid Chelicorophium curvispinum.
3. Systematically increasing predator-free space alters the functional responses (the relationship between prey density and consumption rate) of the amphipod predators by reducing attack rates and lengthening handling times. Crucially, functional response shape also changes subtly from destabilizing Type II towards stabilizing Type III, such that small increases in predator-free space to result in significant reductions in prey consumption at low prey densities.
4. Habitats with superficially similar structural complexity can have considerably divergent consequences for prey population stability in general and, particularly, for invasive prey establishing at low densities in novel habitats.

Eaten alive: cannibalism is enhanced by parasites.

Cannibalism is ubiquitous in nature and especially pervasive in consumers with stage-specific resource utilization in resource-limited environments. Cannibalism is thus influential in the structure and functioning of biological communities. Parasites are also pervasive in nature and, we hypothesize, might affect cannibalism since infection can alter host foraging behaviour. We investigated the effects of a common parasite, the microsporidian Pleistophora mulleri, on the cannibalism rate of its host, the freshwater amphipod Gammarus duebeni celticus. Parasitic infection increased the rate of cannibalism by adults towards uninfected juvenile conspecifics, as measured by adult functional responses, that is, the rate of resource uptake as a function of resource density. This may reflect the increased metabolic requirements of the host as driven by the parasite. Furthermore, when presented with a choice, uninfected adults preferred to cannibalize uninfected rather than infected juvenile conspecifics, probably reflecting selection pressure to avoid the risk of parasite acquisition. By contrast, infected adults were indiscriminate with respect to infection status of their victims, probably owing to metabolic costs of infection and the lack of risk as the cannibals were already infected. Thus parasitism, by enhancing cannibalism rates, may have previously unrecognized effects on stage structure and population dynamics for cannibalistic species and may also act as a selective pressure leading to changes in resource use.

Interchange of entire communities: microbial community coalescence

Microbial communities are enigmatically diverse. We propose a novel view of processes likely affecting microbial assemblages, which could be viewed as the Great American Interchange en miniature: the wholesale exchange among microbial communities resulting from moving pieces of the environment containing entire assemblages. Incidental evidence for such ‘community coalescence’ is accumulating, but such processes are rarely studied, likely because of the absence of suitable terminology or a conceptual framework. We provide the nucleus for such a conceptual foundation for the study of community coalescence, examining factors shaping these events, links to bodies of ecological theory, and we suggest modeling approaches for understanding coalescent communities. We argue for the systematic study of community coalescence because of important functional and applied consequences.

A biological survey method applied to seafloor massive sulphides (SMS) with contagiously distributed hydrothermal-vent fauna

Strategies for mitigation of seafloor massive sulphide (SMS) extraction in the deep sea include establishment of suitable reference sites that allow for studies of natural environmental variability and that can serve as sources of larvae for re-colonisation of extracted hydrothermal fields. In this study, we characterize deep-sea vent communities in Manus Basin (Bismarck Sea, Papua New Guinea) and use macrofaunal data sets from a proposed reference site (South Su) and a proposed mine site (Solwara 1) to test the hypothesis that there was no difference in macrofaunal community structure between the sites. We used dispersion weighting to adjust taxa-abundance matrices to down-weight the contribution of contagious distributions of numerically abundant taxa. Faunal assemblages of 3 habitat types defined by biogenic taxa (2 provannid snails, Alviniconcha spp. and Ifremeria nautilei; and a sessile barnacle, Eochionelasmus ohtai) were distinct from one another and from the vent peripheral assemblage, but were not differentiable from mound-to-mound within a site or between sites. Mussel and tubeworm populations at South Su but not at Solwara 1 enhance the taxonomic and habitat diversity of the proposed reference site. © Inter-Research 2012.

Soil microbes and community coalescence

Community coalescence is a recently introduced term describing the interaction of entire communities and their environments. We here explicitly place the concept of community coalescence in a soil microbial context, exploring intrinsic and extrinsic drivers of such coalescence events. Examples of intrinsic events include the action of earthworms and the dynamics of soil aggregates, while extrinsic events are exemplified by tillage, flooding, litter-fall, outplanting, and the addition of materials containing microbial communities. Aspects of global change may alter the frequency or severity of coalescence events. We highlight functional consequences of community coalescence in soil, and suggest ways to experimentally tackle this phenomenon. Soil ecology as a whole stands to benefit from conceptualizing soil biodiversity in terms of dynamic coalescent microbial assemblages.