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1.  The effect of turbidity on recognition and generalization of predators and non-predators in aquatic ecosystems 
Ecology and Evolution  2012;3(2):268-277.
Recent anthropogenic activities have caused a considerable change in the turbidity of freshwater and marine ecosystems. Concomitant with such perturbations are changes in community composition. Understanding the mechanisms through which species interactions are influenced by anthropogenic change has come to the forefront of many ecological disciplines. Here, we examine how a change in the availability of visual information influences the behavior of prey fish exposed to potential predators and non-predators. When fathead minnows, Pimephales promelas, were conditioned to recognize predators and non-predators in clear water, they showed a highly sophisticated ability to distinguish predators from non-predators. However, when learning occurred under conditions of increased turbidity, the ability of the prey to learn and generalize recognition of predators and non-predators was severely impaired. Our work highlights that changes at the community level associated with anthropogenic perturbations may be mediated through altered trophic interactions, and highlights the need to closely examine behavioral interactions to understand how species interactions change.
doi:10.1002/ece3.454
PMCID: PMC3586637  PMID: 23467451
Anthropogenic change; anti-predator behavior; fathead minnows; predator recognition; risk assessment; turbidity
2.  Issue Information 
Ecology and Evolution  2013;3(2):i-ii.
doi:10.1002/ece3.502
PMCID: PMC3586629  PMID: 23463851
3.  The bacterial parasite Pasteuria ramosa is not killed if it fails to infect: implications for coevolution 
Ecology and Evolution  2012;3(2):197-203.
Strong selection on parasites, as well as on hosts, is crucial for fueling coevolutionary dynamics. Selection will be especially strong if parasites that encounter resistant hosts are destroyed and diluted from the local environment. We tested whether spores of the bacterial parasite Pasteuria ramosa were passed through the gut (the route of infection) of their host, Daphnia magna, and whether passaged spores remained viable for a “second chance” at infecting a new host. In particular, we tested if this viability (estimated via infectivity) depended on host genotype, whether or not the genotype was susceptible, and on initial parasite dose. Our results show that Pasteuria spores generally remain viable after passage through both susceptible and resistant Daphnia. Furthermore, these spores remained infectious even after being frozen for several weeks. If parasites can get a second chance at infecting hosts in the wild, selection for infection success in the first instance will be reduced. This could also weaken reciprocal selection on hosts and slow the coevolutionary process.
doi:10.1002/ece3.438
PMCID: PMC3586630  PMID: 23467806
Daphnia; dilution effect; host–parasite coevolution; Pasteuria
4.  Correlated responses to clonal selection in populations of Daphnia pulicaria: mechanisms of genetic correlation and the creative power of sex 
Ecology and Evolution  2012;3(2):204-216.
Genetic correlations among traits alter evolutionary trajectories due to indirect selection. Pleiotropy, chance linkage, and selection can all lead to genetic correlations, but have different consequences for phenotypic evolution. We sought to assess the mechanisms contributing to correlations with size at maturity in the cyclic parthenogen Daphnia pulicaria. We selected on size in each of four populations that differ in the frequency of sex, and evaluated correlated responses in a life table. Size at advanced adulthood, reproductive output, and adult growth rate clearly showed greater responses in high-sex populations, with a similar pattern in neonate size and r. This pattern is expected only when trait correlations are favored by selection and the frequency of sex favors the creation and demographic expansion of highly fit clones. Juvenile growth and age at maturity did not diverge consistently. The inter-clutch interval appeared to respond more strongly in low-sex populations, but this was not statistically significant. Our data support the hypothesis that correlated selection is the strongest driver of genetic correlations, and suggest that in organisms with both sexual and asexual reproduction, adaptation can be enhanced by recombination.
doi:10.1002/ece3.444
PMCID: PMC3586631  PMID: 23467851
Body size; complex traits; life history; linkage; pleiotropy; recombination
5.  Phylogeography of the Indo-West Pacific maskrays (Dasyatidae, Neotrygon): a complex example of chondrichthyan radiation in the Cenozoic 
Ecology and Evolution  2012;3(2):217-232.
Maskrays of the genus Neotrygon (Dasyatidae) have dispersed widely in the Indo-West Pacific being represented largely by an assemblage of narrow-ranging coastal endemics. Phylogenetic reconstruction methods reproduced nearly identical and statistically robust topologies supporting the monophyly of the genus Neotrygon within the family Dasyatidae, the genus Taeniura being consistently basal to Neotrygon, and Dasyatis being polyphyletic to the genera Taeniurops and Pteroplatytrygon. The Neotrygon kuhlii complex, once considered to be an assemblage of color variants of the same biological species, is the most derived and widely dispersed subgroup of the genus. Mitochondrial (COI, 16S) and nuclear (RAG1) phylogenies used in synergy with molecular dating identified paleoclimatic fluctuations responsible for periods of vicariance and dispersal promoting population fragmentation and speciation in Neotrygon. Signatures of population differentiation exist in N. ningalooensis and N. annotata, yet a large-scale geological event, such as the collision between the Australian and Eurasian Plates, coupled with subsequent sea-level falls, appears to have separated a once homogeneous population of the ancestral form of N. kuhlii into southern Indian Ocean and northern Pacific taxa some 4–16 million years ago. Repeated climatic oscillations, and the subsequent establishment of land and shallow sea connections within and between Australia and parts of the Indo-Malay Archipelago, have both promoted speciation and established zones of secondary contact within the Indian and Pacific Ocean basins.
doi:10.1002/ece3.448
PMCID: PMC3586632  PMID: 23467194
Biodiversity hotspot; cryptic species; marine speciation; maskray; Neotrygon; phylogeography
6.  Extinction hazards in experimental Daphnia magna populations: effects of genotype diversity and environmental variation 
Ecology and Evolution  2012;3(2):233-243.
Extinction is ubiquitous in natural systems and the ultimate fate of all biological populations. However, the factors that contribute to population extinction are still poorly understood, particularly genetic diversity and composition. A laboratory experiment was conducted to examine the influences of environmental variation and genotype diversity on persistence in experimental Daphnia magna populations. Populations were initiated in two blocks with one, two, three, or six randomly selected and equally represented genotypes, fed and checked for extinction daily, and censused twice weekly over a period of 170 days. Our results show no evidence for an effect of the number of genotypes in a population on extinction hazard. Environmental variation had a strong effect on hazards in both experimental blocks, but the direction of the effect differed between blocks. In the first block, variable environments hastened extinction, while in the second block, hazards were reduced under variable food input. This occurred despite greater fluctuations in population size in variable environments in the second block of our experiment. Our results conflict with previous studies, where environmental variation consistently increased extinction risk. They are also at odds with previous studies in other systems that documented significant effects of genetic diversity on population persistence. We speculate that the lack of sexual reproduction, or the phenotypic similarity among our experimental lines, might underlie the lack of a significant effect of genotype diversity in our study.
doi:10.1002/ece3.449
PMCID: PMC3586633  PMID: 23467276
Daphnia magna; environmental variation; extinction; genetic diversity
7.  Plant quality and local adaptation undermine relocation in a bog specialist butterfly 
Ecology and Evolution  2012;3(2):244-254.
The butterfly Boloria aquilonaris is a specialist of oligotrophic ecosystems. Population viability analysis predicted the species to be stable in Belgium and to collapse in the Netherlands with reduced host plant quality expected to drive species decline in the latter. We tested this hypothesis by rearing B. aquilonaris caterpillars from Belgian and Dutch sites on host plants (the cranberry, Vaccinium oxycoccos). Dutch plant quality was lower than Belgian one conferring lower caterpillar growth rate and survival. Reintroduction and/or supplementation may be necessary to ensure the viability of the species in the Netherlands, but some traits may have been selected solely in Dutch caterpillars to cope with gradual changes in host plant quality. To test this hypothesis, the performance of Belgian and Dutch caterpillars fed with plants from both countries were compared. Dutch caterpillars performed well on both plant qualities, whereas Belgian caterpillars could not switch to lower quality plants. This can be considered as an environmentally induced plastic response of caterpillars and/or a local adaptation to plant quality, which precludes the use of Belgian individuals as a unique solution for strengthening Dutch populations. More generally, these results stress that the relevance of local adaptation in selecting source populations for relocation may be as important as restoring habitat quality.
doi:10.1002/ece3.427
PMCID: PMC3586634  PMID: 23467336
Applied ecology; conservation; evolutionary ecology; insect-plant interactions; local adaptation; relocation
8.  Patterns of sexual dimorphism in Mexican alligator lizards, Barisia imbricata 
Ecology and Evolution  2012;3(2):255-261.
We compare morphological characteristics of male and female Barisia imbricata, Mexican alligator lizards, and find that mass, head length, coloration, incidence of scars from conspecifics, tail loss, and frequency of bearing the color/pattern of the opposite sex are all sexually dimorphic traits. Overall size (measured as snout–vent length), on the other hand, is not different between the two sexes. We use data on bite scar frequency and fecundity to evaluate competing hypotheses regarding the selective forces driving these patterns. We contend that sexual selection, acting through male-male competition, may favor larger mass and head size in males, whereas large females are likely favored by natural selection for greater fecundity. In addition, the frequency of opposite-sex patterning in males versus females may indicate that the costs of agonistic interactions among males are severe enough to allow for an alternative mating strategy. Finally, we discuss how sexual and natural selective forces may interact to drive or mask the evolution of sexually dimorphic traits.
doi:10.1002/ece3.455
PMCID: PMC3586635  PMID: 23467394
Body size; Mexican alligator lizards; natural selection; sexual dimorphism; sexual selection
9.  Foliar δ13C response patterns along a moisture gradient arising from genetic variation and phenotypic plasticity in grassland species of Inner Mongolia 
Ecology and Evolution  2012;3(2):262-267.
Plants depend upon both genetic differences and phenotypic plasticity to cope with environmental variation over different timescales. The spatial variation in foliar δ13C levels along a moisture gradient represents an overlay of genetic and plastic responses. We hypothesized that such a spatial variation would be more obvious than the variation arising purely from a plastic response to moisture change. Leymus chinensis and Stipa spp. were sampled from Inner Mongolia along a dry-wet transect, and some of these species were transplanted to an area with a moisture gradient. For Stipa spp., the slope of foliar δ13C and mean annual precipitation along the transect was significantly steeper than that of foliar δ13C and mean annual precipitation after the watering treatment. For L. chinensis, there was a general decreasing trend in foliar δ13C under the different (increasing) watering levels; however, its populations showed an irregular relationship between foliar δ13C and moisture origin. Therefore, support for our hypothesis was obtained from Stipa spp., but not from L. chinensis.
doi:10.1002/ece3.453
PMCID: PMC3586636  PMID: 23467429
Genetic; Leymus chinensis; plastic; Stipa grandis; WUE; δ13C
10.  Unique inhibitory cascade pattern of molars in canids contributing to their potential to evolutionary plasticity of diet 
Ecology and Evolution  2013;3(2):278-285.
Developmental origins that guide the evolution of dental morphology and dental formulae are fundamental subjects in mammalian evolution. In a previous study, a developmental model termed the inhibitory cascade model was established. This model could explain variations in relative molar sizes and loss of the lower third molars, which sometimes reflect diet, in murine rodents and other mammals. Here, I investigated the pattern of relative molar sizes (inhibitory cascade pattern) in canids, a taxon exhibiting a wide range of dietary habits. I found that interspecific variation in canid molars suggests a unique inhibitory cascade pattern that differs from that in murine rodents and other previously reported mammals, and that this variation reflects dietary habits. This unique variability in molars was also observed in individual variation in canid species. According to these observations, canid species have greater variability in the relative sizes of first molars (carnassials), which are functionally important for dietary adaptation in the Carnivora. In conclusion, an inhibitory cascade that differs from that in murine rodents and other mammals may have contributed to diverse dietary patterns and to their parallel evolution in canids.
doi:10.1002/ece3.436
PMCID: PMC3586638  PMID: 23467478
Canidae; Carnivora; dental formulae; dental morphology; evolvability; inhibitory cascade
11.  Cultural legacies, fire ecology, and environmental change in the Stone Country of Arnhem Land and Kakadu National Park, Australia 
Ecology and Evolution  2013;3(2):286-297.
We use the fire ecology and biogeographical patterns of Callitris intratropica, a fire-sensitive conifer, and the Asian water buffalo (Bubalus bubalis), an introduced mega-herbivore, to examine the hypothesis that the continuation of Aboriginal burning and cultural integration of buffalo contribute to greater savanna heterogeneity and diversity in central Arnhem Land (CAL) than Kakadu National Park (KNP). The ‘Stone Country’ of the Arnhem Plateau, extending from KNP to CAL, is a globally renowned social–ecological system, managed for millennia by Bininj-Kunwok Aboriginal clans. Regional species declines have been attributed to the cessation of patchy burning by Aborigines. Whereas the KNP Stone Country is a modern wilderness, managed through prescribed burning and buffalo eradication, CAL remains a stronghold for Aboriginal management where buffalo have been culturally integrated. We surveyed the plant community and the presence of buffalo tracks among intact and fire-damaged C. intratropica groves and the savanna matrix in KNP and CAL. Aerial surveys of C. intratropica grove condition were used to examine the composition of savanna vegetation across the Stone Country. The plant community in intact C. intratropica groves had higher stem counts of shrubs and small trees and higher proportions of fire-sensitive plant species than degraded groves and the savanna matrix. A higher proportion of intact C. intratropica groves in CAL therefore indicated greater gamma diversity and habitat heterogeneity than the KNP Stone Country. Interactions among buffalo, fire, and C. intratropica suggested that buffalo also contributed to these patterns. Our results suggest linkages between ecological and cultural integrity at broad spatial scales across a complex landscape. Buffalo may provide a tool for mitigating destructive fires; however, their interactions require further study. Sustainability in the Stone Country depends upon adaptive management that rehabilitates the coupling of indigenous culture, disturbance, and natural resources.
doi:10.1002/ece3.460
PMCID: PMC3586639  PMID: 23467505
Aboriginal landscape burning; Coupled human natural systems; fire ecology; fire management; habitat heterogeneity; landscape history; plant community diversity; tropical savanna
12.  Molecular and phylogenetic characterization of honey bee viruses, Nosema microsporidia, protozoan parasites, and parasitic mites in China 
Ecology and Evolution  2013;3(2):298-311.
China has the largest number of managed honey bee colonies, which produce the highest quantity of honey and royal jelly in the world; however, the presence of honey bee pathogens and parasites has never been rigorously identified in Chinese apiaries. We thus conducted a molecular survey of honey bee RNA viruses, Nosema microsporidia, protozoan parasites, and tracheal mites associated with nonnative Apis mellifera ligustica and native Apis cerana cerana colonies in China. We found the presence of black queen cell virus (BQCV), chronic bee paralysis virus (CBPV), deformed wing virus (DWV), Israeli acute paralysis virus (IAPV), and sacbrood virus (SBV), but not that of acute bee paralysis virus (ABPV) or Kashmir bee virus (KBV). DWV was the most prevalent in the tested samples. Phylogenies of Chinese viral isolates demonstrated that genetically heterogeneous populations of BQCV, CBPV, DWV, and A. cerana-infecting SBV, and relatively homogenous populations of IAPV and A. meliifera-infecting new strain of SBV with single origins, are spread in Chinese apiaries. Similar to previous observations in many countries, Nosema ceranae, but not Nosema apis, was prevalent in the tested samples. Crithidia mellificae, but not Apicystis bombi was found in five samples, including one A. c. cerana colony, demonstrating that C. mellificae is capable of infecting multiple honey bee species. Based on kinetoplast-encoded cytochrome b sequences, the C. mellificae isolate from A. c. cerana represents a novel haplotype with 19 nucleotide differences from the Chinese and Japanese isolates from A. m. ligustica. This suggests that A. c. cerana is the native host for this specific haplotype. The tracheal mite, Acarapis woodi, was detected in one A. m. ligustica colony. Our results demonstrate that honey bee RNA viruses, N. ceranae, C. mellificae, and tracheal mites are present in Chinese apiaries, and some might be originated from native Asian honey bees.
doi:10.1002/ece3.464
PMCID: PMC3586640  PMID: 23467539
Chinese apiculture; honey bee pathogens and parasites; native Asian honey bees; nonnative European honey bees
13.  Genetic diversity and recombination in natural populations of the nematode-trapping fungus Arthrobotrys oligospora from China 
Ecology and Evolution  2013;3(2):312-325.
Nematophagous fungi can trap and capture nematodes and other small invertebrates. This unique ability has made them ideal organisms from which to develop biological control agents against plant- and animal-parasitic nematodes. However, effective application of biocontrol agents in the field requires a comprehensive understanding about the ecology and population genetics of the nematophagous fungi in natural environments. Here, we genotyped 228 strains of the nematode-trapping fungus Arthrobotrys oligospora using 12 single nucleotide polymorphic markers located on eight random DNA fragments. The strains were from different ecological niches and geographical regions from China. Our analyses identified that ecological niche separations contributed significantly, whereas geographic separation contributed relatively little to the overall genetic variation in our samples of A. oligospora. Interestingly, populations from stressful environments seemed to be more variable and showed more evidence for recombination than those from benign environments at the same geographic areas. We discussed the implications of our results to the conservation and biocontrol application of A. oligospora in agriculture and forestry.
doi:10.1002/ece3.450
PMCID: PMC3586641  PMID: 23467563
Ecological genetics; fungi; population ecology; population genetics – empirical
14.  Effects of predation environment and food availability on somatic growth in the Livebearing Fish Brachyrhaphis rhabdophora (Pisces: Poeciliidae) 
Ecology and Evolution  2013;3(2):326-333.
Variation in somatic growth rates is of great interest to biologists because of the relationship between growth and other fitness-determining traits, and it results from both genetic and environmentally induced variation (i.e. plasticity). Theoretical predictions suggest that mean somatic growth rates and the shape of the reaction norm for growth can be influenced by variation in predator-induced mortality rates. Few studies have focused on variation in reaction norms for growth in response to resource availability between high-predation and low-predation environments. We used juvenile Brachyrhaphis rhabdophora from high-predation and low-predation environments to test for variation in mean growth rates and for variation in reaction norms for growth at two levels of food availability in a common-environment experiment. To test for variation in growth rates in the field, we compared somatic growth rates in juveniles in high-predation and low-predation environments. In the common-environment experiment, mean growth rates did not differ between fish from differing predation environments, but the interaction between predation environment and food level took the form of a crossing reaction norm for both growth in length and mass. Fish from low-predation environments exhibited no significant difference in growth rate between high and low food treatments. In contrast, fish from high-predation environments exhibited variation in growth rates between high and low food treatments, with higher food availability resulting in higher growth rates. In the field, individuals in the high-predation environment grow at a faster rate than those in low-predation environments at the smallest sizes (comparable to sizes in the common-environment experiment). These data provide no evidence for evolved differences in mean growth rates between predation environments. However, fish from high-predation environments exhibited greater plasticity in growth rates in response to resource availability suggesting that predation environments may exhibit increased variation in food availability for prey fish and consequent selection for plasticity.
doi:10.1002/ece3.459
PMCID: PMC3586642  PMID: 23467582
Brachyrhaphis rhabdophora; crossing reaction norm; food availability; growth rate; predation
15.  Divergence in genital morphology may contribute to mechanical reproductive isolation in a millipede 
Ecology and Evolution  2013;3(2):334-343.
Genitalia appear to evolve rapidly and divergently in taxa with internal fertilization. The current consensus is that intense directional sexual selection drives the rapid evolution of genitalia. Recent research on the millipede Antichiropus variabilis suggests that the male genitalia are currently experiencing stabilizing selection – a pattern of selection expected for lock-and-key structures that enforce mate recognition and reproductive isolation. Here, we investigate how divergence in genital morphology affects reproductive compatibility among isolated populations of A. variabilis. Females from a focal population were mated first to a male from their own population and, second, to a male from one of two populations with divergent genital morphology. We observed variation in mating behavior that might indicate the emergence of precopulatory reproductive barriers: males from one divergent population took significantly longer to recognize females and exhibited mechanical difficulty in genital insertion. Moreover, we observed very low paternity success for extra-population males who were successful in copulating. Our data suggest that divergence in genital shape may be contributing to reproductive isolation, and incipient speciation among isolated populations of A. variabilis.
doi:10.1002/ece3.466
PMCID: PMC3586643  PMID: 23467632
Lock-and-key; male genitalia; Millipede; population cross; sexual selection; species mate recognition
16.  Environmental heterogeneity, dispersal mode, and co-occurrence in stream macroinvertebrates 
Ecology and Evolution  2013;3(2):344-355.
Both environmental heterogeneity and mode of dispersal may affect species co-occurrence in metacommunities. Aquatic invertebrates were sampled in 20–30 streams in each of three drainage basins, differing considerably in environmental heterogeneity. Each drainage basin was further divided into two equally sized sets of sites, again differing profoundly in environmental heterogeneity. Benthic invertebrate data were divided into three groups of taxa based on overland dispersal modes: passive dispersers with aquatic adults, passive dispersers with terrestrial winged adults, and active dispersers with terrestrial winged adults. The co-occurrence of taxa in each dispersal mode group, drainage basin, and heterogeneity site subset was measured using the C-score and its standardized effect size. The probability of finding high levels of species segregation tended to increase with environmental heterogeneity across the drainage basins. These patterns were, however, contingent on both dispersal mode and drainage basin. It thus appears that environmental heterogeneity and dispersal mode interact in affecting co-occurrence in metacommunities, with passive dispersers with aquatic adults showing random patterns irrespective of environmental heterogeneity, and active dispersers with terrestrial winged adults showing increasing segregation with increasing environmental heterogeneity.
doi:10.1002/ece3.470
PMCID: PMC3586644  PMID: 23467653
Co-occurrence; dispersal; environmental heterogeneity; headwater streams; metacommunities
17.  Relative performance of hybrid nestlings in Ficedula flycatchers: a translocation experiment 
Ecology and Evolution  2013;3(2):356-364.
Ecological speciation predicts that hybrids should experience relatively low fitness in the local environments of their parental species. In this study, we performed a translocation experiment of nestling hybrids between collared and pied flycatchers into the nests of conspecific pairs of their parental species. Our aim was to compare the performance of hybrids with purebred nestlings. Nestling collared flycatchers are known to beg and grow faster than nestling pied flycatchers under favorable conditions, but to experience higher mortality than nestling pied flycatchers under food limitation. The experiment was performed relatively late in the breeding season when food is limited. If hybrid nestlings have an intermediate growth potential and begging intensity, we expected them to beg and grow faster, but also to experience lower survival than pied flycatchers. In comparison with nestling collared flycatchers, we expected them to beg and grow slower, but to survive better. We found that nestling collared flycatchers indeed begged significantly faster and experienced higher mortality than nestling hybrids. Moreover, nestling hybrids had higher weight and tended to beg faster than nestling pied flycatchers, but we did not detect a difference in survival between the latter two groups of nestlings. We conclude that hybrid Ficedula nestlings appear to have a better intrinsic adaptation to food limitation late in the breeding season compared with nestling collared flycatchers. We discuss possible implications for gene flow between the two species.
doi:10.1002/ece3.472
PMCID: PMC3586645  PMID: 23467681
Environmentally dependent hybrid fitness; genetic incompatibility; hybridization; life history; post-zygotic isolation; speciation
18.  Divergence of premating behaviors in the closely related species Drosophila subquinaria and D. recens 
Ecology and Evolution  2013;3(2):365-374.
Most animal species use distinctive courship patterns to choose among potential mates. Over time, the sensory signaling and preferences used during courtship can diverge among groups that are reproductively isolated. This divergence of signal traits and preferences is thought to be an important cause of behavioral isolation during the speciation process. Here, we examine the sensory modalities used in courtship by two closely related species, Drosophila subquinaria and Drosophila recens, which overlap in geographic range and are incompletely reproductively isolated. We use observational studies of courtship patterns and manipulation of male and female sensory modalities to determine the relative roles of visual, olfactory, gustatory, and auditory signals during conspecific mate choice. We find that sex-specific, species-specific, and population-specific cues are used during mate acquisition within populations of D. subquinaria and D. recens. We identify shifts in both male and female sensory modalities between species, and also between populations of D. subquinaria. Our results indicate that divergence in mating signals and preferences have occurred on a relatively short timescale within and between these species. Finally, we suggest that because olfactory cues are essential for D. subquinaria females to mate within species, they may also underlie variation in behavioral discrimination across populations and species.
doi:10.1002/ece3.477
PMCID: PMC3586646  PMID: 23467696
Mate choice; olfaction; premating isolation; reproductive isolation
19.  Diurnal patterns of gas-exchange and metabolic pools in tundra plants during three phases of the arctic growing season 
Ecology and Evolution  2013;3(2):375-388.
Arctic tundra plant communities are subject to a short growing season that is the primary period in which carbon is sequestered for growth and survival. This period is often characterized by 24-h photoperiods for several months a year. To compensate for the short growing season tundra plants may extend their carbon uptake capacity on a diurnal basis, but whether this is true remains unknown. Here, we examined in situ diurnal patterns of physiological activity and foliar metabolites during the early, mid, and late growing season in seven arctic species under light-saturated conditions. We found clear diurnal patterns in photosynthesis and respiration, with midday peaks and midnight lulls indicative of circadian regulation. Diurnal patterns in foliar metabolite concentrations were less distinct between the species and across seasons, suggesting that metabolic pools are likely governed by proximate external factors. This understanding of diurnal physiology will also enhance the parameterization of process-based models, which will aid in better predicting future carbon dynamics for the tundra. This becomes even more critical considering the rapid changes that are occurring circumpolarly that are altering plant community structure, function, and ultimately regional and global carbon budgets.
doi:10.1002/ece3.467
PMCID: PMC3586647  PMID: 23467719
Alaska; circadian clock; photoperiod; photosynthesis; respiration; sugars; total nonstructural carbohydrates; tussock tundra
20.  Seasonal dimorphism in the horny bills of sparrows 
Ecology and Evolution  2013;3(2):389-398.
Bill size is often viewed as a species-specific adaptation for feeding, but it sometimes varies between sexes, suggesting that sexual selection or intersexual competition may also be important. Hypotheses to explain sexual dimorphism in avian bill size include divergence in feeding niche or thermoregulatory demands, intrasexual selection based on increased competition among males, or female preference. Birds also show seasonal changes in bill size due to shifts in the balance between growth rate and wear, which may be due to diet or endogenous rhythms in growth. Insight into the function of dimorphism can be gained using the novel approach of digital x-ray imaging of museum skins to examine the degree to which the skeletal core or the rhamphotheca contribute to overall dimorphism. The rhamphotheca is ever-growing and ever-wearing, varying in size throughout life; whereas the skeletal core shows determinant growth. Because tidal marsh sparrows are more dimorphic in bill size than related taxa, we selected two marsh taxa to investigate dimorphism and seasonality in the size of the overall bill, the skeletal core, and the rhamphotheca. Bill size varied by sex and season, with males having larger bills than females, and bill size increasing from nonbreeding to breeding season more in males. Skeletal bill size varied with season, but not sex. The rhamphotheca varied primarily with sex; males had a larger rhamphotheca (corrected for skeletal bill size), which showed a greater seasonal increase than females. The rhamphotheca, rather than the skeletal bill, was responsible for sexual dimorphism in overall bill size, which was particularly well developed in the breeding season. The size of the rhamphotheca may be a condition-based character that is shaped by sexual selection. These results are consistent with the evidence that bill size is influenced by sexual selection as well as trophic ecology.
doi:10.1002/ece3.474
PMCID: PMC3586648  PMID: 23467758
Bill size; bird beaks; emberizidae; rhamphotheca; salt marsh birds; sexual selection; tidal marsh birds
21.  Epigenetic regulation of adaptive responses of forest tree species to the environment 
Ecology and Evolution  2013;3(2):399-415.
Epigenetic variation is likely to contribute to the phenotypic plasticity and adaptative capacity of plant species, and may be especially important for long-lived organisms with complex life cycles, including forest trees. Diverse environmental stresses and hybridization/polyploidization events can create reversible heritable epigenetic marks that can be transmitted to subsequent generations as a form of molecular “memory”. Epigenetic changes might also contribute to the ability of plants to colonize or persist in variable environments. In this review, we provide an overview of recent data on epigenetic mechanisms involved in developmental processes and responses to environmental cues in plant, with a focus on forest tree species. We consider the possible role of forest tree epigenetics as a new source of adaptive traits in plant breeding, biotechnology, and ecosystem conservation under rapid climate change.
doi:10.1002/ece3.461
PMCID: PMC3586649  PMID: 23467802
Adaptive response; environmental stress; epigenetic memory of stressful conditions; epigenetics; forest trees; phenotypic plasticity
22.  Simulating evolutionary responses of an introgressed insect resistance trait for ecological effect assessment of transgene flow: a model for supporting informed decision-making in environmental risk assessment 
Ecology and Evolution  2013;3(2):416-423.
Predicting outcomes of transgene flow from arable crops requires a system perspective that considers ecological and evolutionary processes within a landscape context. In Europe, the arable weed Raphanus raphanistrum is a potential hybridization partner of oilseed rape, and the two species are ecologically linked through the common herbivores Meligethes spp. Observations in Switzerland show that high densities of Meligethes beetles maintained by oilseed rape crops can lead to considerable damage on R. raphanistrum. We asked how increased insect resistance in R. raphanistrum – as might be acquired through introgression from transgenic oilseed rape – would affect seed production under natural herbivore pressure. In simulation experiments, plants protected against Meligethes beetles produced about twice as many seeds as unprotected plants. All stages in the development of reproductive structures from buds to pods were negatively affected by the herbivore, with the transition from buds to flowers being the most vulnerable. We conclude that resistance to Meligethes beetles could confer a considerable selective advantage upon R. raphanistrum in regions where oilseed rape is widely grown.
doi:10.1002/ece3.463
PMCID: PMC3586650  PMID: 23467842
Apparent competition; crop–wild gene flow; Meligethes beetles; oilseed rape; Raphanus raphanistrum; transgenic plants
23.  Adaptive responses and invasion: the role of plasticity and evolution in snail shell morphology 
Ecology and Evolution  2013;3(2):424-436.
Invasive species often exhibit either evolved or plastic adaptations in response to spatially varying environmental conditions. We investigated whether evolved or plastic adaptation was driving variation in shell morphology among invasive populations of the New Zealand mud snail (Potamopyrgus antipodarum) in the western United States. We found that invasive populations exhibit considerable shell shape variation and inhabit a variety of flow velocity habitats. We investigated the importance of evolution and plasticity by examining variation in shell morphological traits 1) between the parental and F1 generations for each population and 2) among populations of the first lab generation (F1) in a common garden, full-sib design using Canonical Variate Analyses (CVA). We compared the F1 generation to the parental lineages and found significant differences in overall shell shape indicating a plastic response. However, when examining differences among the F1 populations, we found that they maintained among-population shell shape differences, indicating a genetic response. The F1 generation exhibited a smaller shell morph more suited to the low-flow common garden environment within a single generation. Our results suggest that phenotypic plasticity in conjunction with evolution may be driving variation in shell morphology of this widespread invasive snail.
doi:10.1002/ece3.471
PMCID: PMC3586651  PMID: 23467920
Adaptive evolution; Canonical variate analysis; morphometric landmarks; phenotypic plasticity; Potamopyrgus antipodarum; shell morphology
24.  Intraspecific variation buffers projected climate change impacts on Pinus contorta 
Ecology and Evolution  2013;3(2):437-449.
Species distribution modeling (SDM) is an important tool to assess the impact of global environmental change. Many species exhibit ecologically relevant intraspecific variation, and few studies have analyzed its relevance for SDM. Here, we compared three SDM techniques for the highly variable species Pinus contorta. First, applying a conventional SDM approach, we used MaxEnt to model the subject as a single species (species model), based on presence–absence observations. Second, we used MaxEnt to model each of the three most prevalent subspecies independently and combined their projected distributions (subspecies model). Finally, we used a universal growth transfer function (UTF), an approach to incorporate intraspecific variation utilizing provenance trial tree growth data. Different model approaches performed similarly when predicting current distributions. MaxEnt model discrimination was greater (AUC – species model: 0.94, subspecies model: 0.95, UTF: 0.89), but the UTF was better calibrated (slope and bias – species model: 1.31 and −0.58, subspecies model: 1.44 and −0.43, UTF: 1.01 and 0.04, respectively). Contrastingly, for future climatic conditions, projections of lodgepole pine habitat suitability diverged. In particular, when the species' intraspecific variability was acknowledged, the species was projected to better tolerate climatic change as related to suitable habitat without migration (subspecies model: 26% habitat loss or UTF: 24% habitat loss vs. species model: 60% habitat loss), and given unlimited migration may increase amount of suitable habitat (subspecies model: 8% habitat gain or UTF: 12% habitat gain vs. species model: 51% habitat loss) in the climatic period 2070–2100 (SRES A2 scenario, HADCM3). We conclude that models derived from within-species data produce different and better projections, and coincide with ecological theory. Furthermore, we conclude that intraspecific variation may buffer against adverse effects of climate change. A key future research challenge lies in assessing the extent to which species can utilize intraspecific variation under rapid environmental change.
doi:10.1002/ece3.426
PMCID: PMC3586652  PMID: 23467191
Intraspecific diversity; lodgepole pine; niche modeling; North America; range shift; within-species variability
25.  Functional endemism: population connectivity, shifting baselines, and the scale of human experience 
Ecology and Evolution  2012;3(2):450-456.
Quantifying population connectivity is important for visualizing the spatial and temporal scales that conservation measures act upon. Traditionally, migration based on genetic data has been reported in migrants per generation. However, the temporal scales over which this migration may occur do not necessarily accommodate the scales over which human perturbations occur, leaving the potential for a disconnect between population genetic data and conservation action based on those data. Here, we present a new metric called the “Rule of Memory”, which helps conservation practitioners to interpret “migrants per generation” in the context both of human modified ecosystems and the cultural memory of those doing the modification. Our rule states that clades should be considered functionally endemic regardless of their actual taxonomic designation if the migration between locations is insufficient to maintain a viable population over the timescales of one human generation (20 years). Since larger animals are more likely to be remembered, we quantify the relationship between migrants per human (N) and body mass of the organism in question (M) with the formula N = 10M−1. We then use the coral reef fish Pomacentrus moluccensis to demonstrate the taxonomic and spatial scales over which this rule can be applied. Going beyond minimum viable population literature, this metric assesses the probability that a clade's existence will be forgotten by people throughout its range during a period of extirpation. Because conservation plans are predicated on having well-established baselines, a loss of a species over the range of one human generation evokes the likelihood of that species no longer being recognized as a member of an ecosystem, and thus being excluded in restoration or conservation prioritization. [Correction added on 26 December 2012, after first online publication: this formula has been corrected to N=10M−1].
doi:10.1002/ece3.446
PMCID: PMC3586653  PMID: 23467269
Coral Reefs; historical ecology; migrants per generation; population connectivity; shifting baselines

Results 1-25 (369)