Parsing parallel evolution:Ecological divergence and differential gene expression in the adaptive radiations of thick-lipped Midas cichlid fishes from Nicaragua

The study of parallel evolution facilitates the discovery of common rules of diversification. Here, we examine the repeated evolution of thick lips in Midas cichlid fishes (the Amphilophus citrinellus species complex) - from two Great Lakes and two crater lakes in Nicaragua - to assess whether similar changes in ecology, phenotypic trophic traits and gene expression accompany parallel trait evolution. Using next-generation sequencing technology, we characterize transcriptome-wide differential gene expression in the lips of wild-caught sympatric thick- and thin-lipped cichlids from all four instances of repeated thick-lip evolution. Six genes (apolipoprotein D, myelin-associated glycoprotein precursor, four-and-a-half LIM domain protein 2, calpain-9, GTPase IMAP family member 8-like and one hypothetical protein) are significantly underexpressed in the thick-lipped morph across all four lakes. However, other aspects of lips' gene expression in sympatric morphs differ in a lake-specific pattern, including the magnitude of differentially expressed genes (97-510). Generally, fewer genes are differentially expressed among morphs in the younger crater lakes than in those from the older Great Lakes. Body shape, lower pharyngeal jaw size and shape, and stable isotopes (dC and dN) differ between all sympatric morphs, with the greatest differentiation in the Great Lake Nicaragua. Some ecological traits evolve in parallel (those related to foraging ecology; e.g. lip size, body and head shape) but others, somewhat surprisingly, do not (those related to diet and food processing; e.g. jaw size and shape, stable isotopes). Taken together, this case of parallelism among thick- and thin-lipped cichlids shows a mosaic pattern of parallel and nonparallel evolution. © 2012 Blackwell Publishing Ltd.

The spatial ecology of the whiskered bat (Myotis mystacinus) at the western extreme of its range provides evidence of regional adaptation

Although widespread, the ecology of the whiskered bat, Myotis mystacinus in Europe remains poorly understood. Ireland is positioned at the most western extreme of this species' range. To ascertain the ecology of M. mystacinus at its geographic range extreme, the roosting behaviour, home range and habitat use of females in a maternity roost in Ireland was investigated by radio-tracking. M. mystacinus were active in a diversity of habitats: namely, mixed woodland, riparian vegetation, arable land and rough grassland. However, only mixed woodland and riparian habitats were selected as core foraging areas. This is in contrast to a previous study from Britain where only pasture was utilised but is in agreement with data from Slovakia, where woodland was also selected, whilst riparian areas were also utilised by this species in Germany. A high degree of overlap in the foraging areas of individuals was observed. A total of seven roosts were utilised by tracked bats and roost switching behaviour was observed. We discuss our contrasting results in respect to range limitations, regional variability in landscape structure and the composition of bat communities. The present results have implications for the conservation of M. mystacinus within Ireland and other parts of its range, highlighting the need for range wide ecological studies. Regional variability in the ecology of bats related to landscape factors is an important consideration for bat conservation and therefore must be incorporated into future management plans. (C) 2012 Deutsche Gesellschaft fur Saugetierkunde. Published by Elsevier GmbH. All rights reserved.

Leatherback turtles satellite-tagged in European waters

The North Atlantic is considered a stronghold for the critically endangered leatherback sea turtle. However, limited information exists regarding the movements of individuals to and from the seas off Europe's northwesterly fringe, an area where featherbacks have been historically sighted for the past 200 yr. Here, we used satellite telemetry to record the movements and behaviour of 2 individuals bycaught in fisheries off the southwest coast of Ireland. The turtle T1 (tagged 1 September 2005; female; tracked 375 d) immediately travelled south via Madeira and the Canaries, before residing in West African waters for 3 mo. In spring, T1 migrated north towards Newfoundland where transmissions ceased. T2 (29 June 2006; male; 233 d) travelled south for a short period before spending 66 d west of the Bay of Biscay, an area previously asserted as a high-use area for leatherbacks. This prolonged high latitude summer residence corresponded with a mesoscale feature evident from satellite imagery, with the implication that this turtle had found a rich feeding site. A marked change in dive behaviour was apparent as the turtle exited this feature and provided useful insights on leatherback diving behaviour. T2 headed south in October 2006, and performed the deepest-ever dive recorded by a reptile (1280 m) southwest of Cape Verde. Unlike T1, T2 swam southwest towards Brazil before approaching the major nesting beaches of French Guiana and Surinam. Importantly, these tracks document the movement of leatherbacks from one of the remotest foraging grounds in the North Atlantic. © Inter-Research 2008.

Behavioural context of multi-scale species distribution models assessed by radio-tracking

Incorporating ecological processes and animal behaviour into Species Distribution Models (SDMs) is difficult. In species with a central resting or breeding place, there can be conflict between the environmental requirements of the 'central place' and foraging habitat. We apply a multi-scale SDM to examine habitat trade-offs between the central place, roost sites, and foraging habitat in . Myotis nattereri. We validate these derived associations using habitat selection from behavioural observations of radio-tracked bats. A Generalised Linear Model (GLM) of roost occurrence using land cover variables with mixed spatial scales indicated roost occurrence was positively associated with woodland on a fine scale and pasture on a broad scale. Habitat selection of radio-tracked bats mirrored the SDM with bats selecting for woodland in the immediate vicinity of individual roosts but avoiding this habitat in foraging areas, whilst pasture was significantly positively selected for in foraging areas. Using habitat selection derived from radio-tracking enables a multi-scale SDM to be interpreted in a behavioural context. We suggest that the multi-scale SDM of . M. nattereri describes a trade-off between the central place and foraging habitat. Multi-scale methods provide a greater understanding of the ecological processes which determine where species occur and allow integration of behavioural processes into SDMs. The findings have implications when assessing the resource use of a species at a single point in time. Doing so could lead to misinterpretation of habitat requirements as these can change within a short time period depending on specific behaviour, particularly if detectability changes depending on behaviour. © 2011 Gesellschaft für ökologie.

A generalized functional response for predators that switch between multiple prey species

We develop a theory for the food intake of a predator that can switch between multiple prey species. The theory addresses empirical observations of prey switching and is based on the behavioural assumption that a predator tends to continue feeding on prey that are similar to the prey it has consumed last, in terms of, e.g., their morphology, defences, location, habitat choice, or behaviour. From a predator's dietary history and the assumed similarity relationship among prey species, we derive a general closed-form multi-species functional response for describing predators switching between multiple prey species. Our theory includes the Holling type II functional response as a special case and makes consistent predictions when populations of equivalent prey are aggregated or split. An analysis of the derived functional response enables us to highlight the following five main findings. (1) Prey switching leads to an approximate power-law relationship between ratios of prey abundance and prey intake, consistent with experimental data. (2) In agreement with empirical observations, the theory predicts an upper limit of 2 for the exponent of such power laws. (3) Our theory predicts deviations from power-law switching at very low and very high prey-abundance ratios. (4) The theory can predict the diet composition of a predator feeding on multiple prey species from diet observations for predators feeding only on pairs of prey species. (5) Predators foraging on more prey species will show less pronounced prey switching than predators foraging on fewer prey species, thus providing a natural explanation for the known difficulties of observing prey switching in the field. (C) 2013 Elsevier Ltd. All rights reserved.

Historical data reveal power-law dispersal patterns of invasive aquatic species

Understanding how invasive species spread is of particular concern in the current era of globalisation and rapid environmental change. The occurrence of super-diffusive movements within the context of Lévy flights has been discussed with respect to particle physics, human movements, microzooplankton, disease spread in global epidemiology and animal foraging behaviour. Super-diffusive movements provide a theoretical explanation for the rapid spread of organisms and disease, but their applicability to empirical data on the historic spread of organisms has rarely been tested. This study focuses on the role of long-distance dispersal in the invasion dynamics of aquatic invasive species across three contrasting areas and spatial scales: open ocean (north-east Atlantic), enclosed sea (Mediterranean) and an island environment (Ireland). Study species included five freshwater plant species, Azolla filiculoides, Elodea canadensis, Lagarosiphon major, Elodea nuttallii and Lemna minuta; and ten species of marine algae, Asparagopsis armata, Antithamnionella elegans, Antithamnionella ternifolia, Codium fragile, Colpomenia peregrina, Caulerpa taxifolia, Dasysiphonia sp., Sargassum muticum, Undaria pinnatifida and Womersleyella setacea. A simulation model is constructed to show the validity of using historical data to reconstruct dispersal kernels. Lévy movement patterns similar to those previously observed in humans and wild animals are evident in the re-constructed dispersal pattern of invasive aquatic species. Such patterns may be widespread among invasive species and could be exacerbated by further development of trade networks, human travel and environmental change. These findings have implications for our ability to predict and manage future invasions, and improve our understanding of the potential for spread of organisms including infectious diseases, plant pests and genetically modified organisms.

Zooarchaeology at the Niah Caves, Sarawak:Context and Research Issues INTRODUCTION

Extensive archaeological excavations in the Niah Caves (Sarawak, Malaysian Borneo) over the past 50 years have produced perhaps 750 000 fragments of vertebrate bone, one of the largest faunal assemblages in the region, This paper introduces a series of research studies examining different aspects of the Niah fauna, and discusses how they are contributing to, and shaping, regional research agendas relating to prehistoric environments and societies in Island Southeast Asia. Zooarchaeology has traditionally had a rather 'Cinderella' status here, but the ongoing programme of study of the Niah Caves fauna is demonstrating the remarkable potential of this material to address questions of Pleistocene and Holocene climate and environment, biodiversity, human activities within caves, people's engagement with the landscapes they inhabited as foragers and farmers, and the nature of the transition from foraging to farming. Copyright (C) 2009 John Wiley & Sons, Ltd.

The 'human revolution' in lowland tropical Southeast Asia:the antiquity and behavior of anatomically modern humans at Niah Cave (Sarawak, Borneo)

Recent research in Europe, Africa, and Southeast Asia suggests that we can no longer assume a direct and exclusive link between anatomically modern humans and behavioral modernity (the 'human revolution'), and assume that the presence of either one implies the presence of the other: discussions of the emergence of cultural complexity have to proceed with greater scrutiny of the evidence on a site-by-site basis to establish secure associations between the archaeology present there and the hominins who created it. This paper presents one such case study: Niah Cave in Sarawak on the island of Borneo, famous for the discovery in 1958 in the West Mouth of the Great Cave of a modern human skull, the 'Deep Skull,' controversially associated with radiocarbon dates of ca. 40,000 years before the present. A new chronostratigraphy has been developed through a re-investigation of the lithostratigraphy left by the earlier excavations, AMS-dating using three different comparative pre-treatments including ABOX of charcoal, and U-series using the Diffusion-Absorption model applied to fragments of bones from the Deep Skull itself. Stratigraphic reasons for earlier uncertainties about the antiquity of the skull are examined, and it is shown not to be an `intrusive' artifact. It was probably excavated from fluvial-pond-desiccation deposits that accumulated episodically in a shallow basin immediately behind the cave entrance lip, in a climate that ranged from times of comparative aridity with complete desiccation, to episodes of greater surface wetness, changes attributed to regional climatic fluctuations. Vegetation outside the cave varied significantly over time, including wet lowland forest, montane forest, savannah, and grassland. The new dates and the lithostratigraphy relate the Deep Skull to evidence of episodes of human activity that range in date from ca. 46,000 to ca. 34,000 years ago. Initial investigations of sediment scorching, pollen, palynomorphs, phytoliths, plant macrofossils, and starch grains recovered from existing exposures, and of vertebrates from the current and the earlier excavations, suggest that human foraging during these times was marked by habitat-tailored hunting technologies, the collection and processing of toxic plants for consumption, and, perhaps, the use of fire at some forest-edges. The Niah evidence demonstrates the sophisticated nature of the subsistence behavior developed by modern humans to exploit the tropical environments that they encountered in Southeast Asia, including rainforest. (c) 2006 Elsevier Ltd. All rights reserved.

The emergence of bone technologies at the end of the pleistocene in Southeast Asia:Regional and evolutionary implications

For many decades Palaeolithic research viewed the development of early modern human behaviour as largely one of progress down a path towards the modernity of the present. The European Palaeolithic sequence the most extensively studied was for a long time the yard-stick against which records from other regions were judged. Recent work undertaken in Africa and increasingly Asia, however, now suggests that the European evidence may tell a story that is more parochial and less universal than previously thought. While tracking developments at the large scale (the grand narrative) remains important, there is growing appreciation that to achieve a comprehensive understanding of human behavioural evolution requires an archaeologically regional perspective to balance this. One of the apparent markers of human modernity that has been sought in the global Palaeolithic record, prompted by finds in the European sequence, is innovation in bonebased technologies. As one step in the process of re-evaluating and contextualizing such innovations, in this article we explore the role of prehistoric bone technologies within the Southeast Asian sequence, where they have at least comparable antiquity to Europe and other parts of Asia. We observe a shift in the technological usage of bone from a minor component to a medium of choice during the second half of the Last Termination and into the Holocene. We suggest that this is consistent with it becoming a focus of the kinds of inventive behaviour demanded of foraging communities as they adapted to the far-reaching environmental and demographic changes that were reshaping this region at that time. This record represents one small element of a much wider, much longerterm adaptive process, which we would argue is not confined to the earliest instances of a particular technology or behaviour, but which forms part of an on-going story of our behavioural evolution. © 2012 The McDonald Institute for Archaeological Research.

Prehistoric foragers and farmers in south-east Asia:Renewed investigations at Niah Cave, Sarawak

The paper describes the initial results from renewed investigations at Niah Cave in Sarawak on the island of Borneo, famous for the discovery in 1958 of the c. 40,000-year old 'Deep Skull'. The archaeological sequences from the West Mouth and the other entrances of the cave complex investigated by Tom and Barbara Harrisson and other researchers have potential implications for three major debates regarding the prehistory of south-east Asia: the timing of initial settlement by anatomically modern humans; the means by which they subsisted in the late Pleistocene and early Holocene; and the timing, nature, and causation of the transition from foraging to farming. The new project is informing on all three debates. The critical importance of the Niah stratigraphies was commonly identified - including by Tom Harrisson himself - as because the site provided a continuous sequence of occupation over the past 40,000 years. The present project indicates that Niah was first used at least 45,000 years ago, and probably earlier; that the subsequent Pleistocene and Holocene occupations were highly variable in intensity and character; and that in some periods, perhaps of significant duration, the caves may have been more or less abandoned. The cultural sequence that is emerging from the new investigations may be more typical of cave use in tropical rainforests in south-east Asia than the Harrisson model.