Does desertification diminish biodiversity? Enhancement of ant diversity by shrub invasion in south‐western USA

The conversion of desert grasslands to shrublands is a long‐standing concern in the south‐western United States, but the effects of this change on native animals defy generalization. Here, I consider evidence that shrub invasion and encroachment, particularly that of honey mesquite (Prosopis glandulosa), has led to increasing ecological dominance and diversity of ants in general, as well as increases in specific native taxa. The effects of shrub invasion on ants were measured at two scales: (1) between Chihuahuan Desert landscapes that vary slightly in temperature and strongly in the dominance of mesquite, and (2) across variation in mesquite density occurring within a generally mesquite‐dominated landscape. Ant richness and numerical dominance was measured at pitfall traps over 2 years and baits were used to assess ecological dominance across different temperatures. The mesquite‐dominated Jornada site harboured four times the number of ant foragers found at the relatively ‘pristine’ Sevilleta site, with several ecologically dominant taxa driving this pattern, especially Dorymyrmex bicolor. Species richness and ecological dominance were also greatest at the Jornada. Within the Jornada landscape, turnover in species composition was related to mesquite density, but local richness and abundance was unrelated to mesquite density. Coupled with the results of previous manipulative experiments and comparative studies, there is support for the notion that ant diversity is not negatively affected by shrub invasion but that several taxa prosper from it. The Jornada is uniquely saturated by dominant ant taxa, perhaps as a consequence of an overall high level of shrub availability that provides a reliable source of carbohydrate‐rich plant exudates. This raises important questions about the relationship between environmental degradation, ecosystem productivity, and animal diversity.     

Does intraspecific competition promote variation? A test via synthesis

Competitive diversification, that is, when increasing intraspecific competition promotes population niche expansion, is commonly invoked in evolutionary studies and currently plays a central role in how we conceptualize the process of adaptive diversification. Despite the frequency with which this idea is cited, the empirical evidence for the process is somewhat limited, and the findings of these studies have yet to be weighed objectively through synthesis. Here, we sought to fill this gap by reviewing the existing literature and collecting the data necessary to assess the evidence for competition as a diversifying force. Additionally, we sought to test a more recent hypothesis, which suggests that competition can act to both promote and inhibit dietary diversification depending on the degree to which a consumer depletes its resources. The surprising result of this synthesis was that increasing competition did not have a mean positive effect on population‐level diet breadth or the degree of individual specialization. Instead, we found that increasing intraspecific competition had a restricting effect on population‐level diet breadth in as many cases as it had a diversifying effect. This wide disparity in the effect of competition on consumer diet variation was negatively related to a metric for consumer resource depletion. Altogether, these findings call into question a long‐standing assumption of basic evolutionary models and lend some support to recent theoretical predictions. Specifically, these findings support the idea that competition is primarily diversifying for species with a small effect (per unit biomass) on their resources and that resource depletion limits the diversifying effect of competition for consumers with larger ecological effects.     

Does cross‐acclimation between drought and freezing stress persist over ecologically relevant time spans? A test using the grass Poa pratensis

• Despite evidence that prior exposure to drought can increase subsequent plant freezing tolerance, few studies have explored such interactions over ecologically relevant time spans. We examined the combined effects of drought and subsequent freezing on tiller growth and leaf sugar concentrations in the grass, Poa pratensis .
• We exposed tillers to no drought (?0.04 MP a), moderate drought (?0.19 MP a) or severe drought (?0.42 MP a) for 3 weeks in summer. Tillers were then frozen in autumn or spring at ?5 °C (frost damage) or at 0 °C (control) for 3 days and harvested after a re‐growth period.
• For shoot growth, there was a significant interaction between drought and autumn freezing, whereby the relative effect of freezing on growth was least for the plants previously exposed to severe drought; however, there was no significant interaction between drought and spring freezing. For root growth, there were no significant interactions between drought and freezing in either season. Leaf sugar concentrations increased significantly with drought intensity, but these effects dissipated within a month, prior to the onset of the autumn freezing treatment.
• Overall, our results suggest that interactions between prior drought and subsequent freezing in P. pratensis may be most relevant in the context of autumn freezing, and despite the important role of soluble sugars in increasing both drought and freezing tolerance in this species, the retention of these compounds after drought stress does not appear to explain the occurrence of drought–frost interactions at ecologically relevant time scales.

     

Does shelter enhance early seedling survival in dry environments? A test with eight Mediterranean species

Question: Do solid‐walled polyethylene tubes and mesh fabric tubes improve the short‐term survival of eight Mediterranean tree and shrub species often used in the restoration of arid environments? Location: We conducted two experimental plantations in degraded field sites in the province of Almería (SE Spain), under arid Mediterranean conditions. Methods: One‐year‐old seedlings of Ceratonia siliqua, Juniperus phoenicea, Olea europaea, Pinus halepensis, Pinus pinaster, Quercus coccifera, Quercus ilex and Tetraclinis articulata were planted either sheltered by one of the above shelter tubes, or by being left unsheltered. Survival was recorded the first growing season after planting, which was a very dry season. Results: Overall, seedling survival ranged from as little as 0% to 24%, and tree shelters consistently enhanced survival in Quercus species only, ranging from 16% in walled shelters to 8% in mesh shelters. Shelters failed to boost survival in the six remaining species. Conclusion: The results of this study suggest that both walled and mesh shelters were mostly ineffective at increasing seedling survival for the Mediterranean species used in this experiment; these species coincide with those used in restoration programs. The use of shelters in restoration programs conducted in arid environments should be reconsidered, while walled shelters might be advisable for Mediterranean Quercus species only. Further research is necessary to develop and assess improved types of shelters for arid environments.     

Does alpine grazing reduce blazing? A landscape test of a widely‐held hypothesis

Abstract ‘Alpine grazing reduces blazing’ is a widely and strongly held view concerning the effects of livestock grazing on fuels, and therefore fire behaviour and impact, in Australia's high country landscapes. As a test of this hypothesis, we examined the patterns of burning across the alpine (treeless) landscapes of the Bogong High Plains in Victoria, following the extensive fires of January 2003. Data were collected from multiple transects, each 3–5 km long, with survey points located randomly at either 50, 200 or 500 m intervals. The transects traversed the major regions of the Bogong High Plains, both grazed and ungrazed. At each point, we recorded whether the point was burnt or unburnt, the vegetation type (closed‐heath, open‐heath, grassland or herbfield), the estimated prefire shrub cover, slope, aspect, and a GPS location. At burnt heathland sites, we recorded the minimum twig diameter (an a posteriori measure of fire severity) in a sample of common shrubs. In total, there were 108 km of transect lines, 419 survey points and 4050 twig measurements, with sample points equally distributed across grazed and ungrazed country. The occurrence of fire (i.e. burnt or unburnt) in grazed and ungrazed areas was analysed by logistic regression; the variation in twig diameters by anova . Approximately half of all points were burnt. There was no statistically significant difference between grazed and ungrazed areas in the proportion of points burnt. Fire occurrence was determined primarily by vegetation type, with the proportion burnt being 0.87 for closed‐heath, 0.59 for open‐heath, and 0.13 for grassland and all snow‐patch herbfield points unburnt. In both closed‐heath and open‐heath, grazing did not significantly lower the severity of fire, as measured by the diameter of burnt twigs. We interpret the lack of a grazing effect in terms of shrub dynamics (little or no grazing effect on long‐term cover of taller shrubs), diet and behaviour of cattle (herbs and dwarf shrubs eaten; tall shrubs not eaten and closed‐heath vegetation generally avoided), and fuel flammability (shrubs more flammable than grass). Whatever effects livestock grazing may have on vegetation cover, and therefore fuels in alpine landscapes, they are likely to be highly localized, with such effects unlikely to translate into landscape‐scale reduction of fire occurrence or severity. The use of livestock grazing in Australian alpine environments as a fire abatement practice is not justified on scientific grounds.     

Does whole‐organism performance constrain resource use? A community test with desert lizards

Discovering the mechanisms by which communities of co‐existing species exist has proven to be one of the greatest challenges for evolutionary ecologists. A recent perspective emphasizes the role of functional traits, such as whole‐organism performance, as key limiting factors in the evolution of communities, yet few studies have examined this possibility. We examine how bite force and morphology influence the ability of ten lizard species in a single community to access insect prey, as defined by prey type and prey hardness. We gathered over 3 years of data from a desert lizard community comprised of ten species and found significant variation among species for bite force and prey hardness, as well as significant differences in performance and niche breadth for each species. In general, higher levels of absolute bite force broadens resource accessibility (sizes of prey), and does not generally result in a reduced ability to access smaller prey. For example, large lizard species that are hard biters can still consume soft prey. On the other hand, small lizard species that are weak biters are more limited in their ability to access hard prey, although the overall decline in resource accessibility is modest. Our findings highlight how functional traits can influence which species can access key resources within a community of similar species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Does facilitating pollinator learning impede deceptive orchid attractiveness? A multi‐approach test of avoidance learning

It has often been proposed that nectarless deceptive orchid species exploit naïve pollinators in search of food before they learn to avoid their flowers, and that intraspecific floral trait polymorphism, often noted in this plant group, could prolong the time needed for learning, thus increasing orchid reproductive success. We tested the importance of avoidance learning in a European deceptive orchid, Anacamptis morio, which has been reported to have a highly variable fragrance bouquet among individuals. We used an indirect approach, i.e. we facilitated pollinators’ ability to learn to avoid A. morio by adding anisaldehyde to selected inflorescences, a scent compound that is easily perceived by the natural pollinators and produced in large quantities by the closely related, nectar producing Anacamptis coriophora, a species that shares pollinator species with A. morio. In a series of three experiments (in artificial arrays, in natural populations and in bumblebee behavioural observations), we consistently found no difference either of reproductive success of or visitation rates to scent‐added versus control inflorescences. We also found that the decrease of reproductive success over time in artificial populations of this deceptive species was not as important as expected. Together, these data suggest that pollinators do not fully learn to avoid deceptive inflorescences, and that pollinator avoidance behaviour alone may explain the lower reproductive success usually found in deceptive orchids. We discuss the possible explanations for this pattern in deceptive orchids, particularly in relation to pollinator cognition and learning abilities. Lastly, in light of our results, the potential for higher average reproductive success in deceptive orchids with high phenotypic variability driven by avoidance learning thus appears to be challenged.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

Do invasive ants respond more strongly to carbohydrate availability than co‐occurring non‐invasive ants? A test along an active Anoplolepis gracilipes invasion front

Invasions by non‐native insects can have important ecological impacts, particularly on island ecosystems. However, the factors that promote the success of invaders relative to co‐occurring non‐invasive species remain unresolved. For invasive ants, access to carbohydrate resources via interactions with both extrafloral nectary‐bearing plants and honeydew‐excreting insects may accelerate the invasion process. A first step towards testing this hypothesis is to determine whether invasive ants respond to variation in the availability of carbohydrate resources, and whether this response differs from that of co‐occurring, non‐invasive ants. We investigated the effect of carbohydrate subsidies on the short‐term foraging and hemipteran‐tending behaviours of the invasive ant Anoplolepis gracilipes (Formicidae) and co‐occurring ant species on an extrafloral nectary‐bearing plant by experimentally manipulating carbohydrate levels and tracking ant recruitment. We conducted experiments in 2 years at two sites: one site was invaded by A. gracilipes prior to 2007 and the other became invaded during the course of our study, allowing pre‐ (2007) and post‐invasion (2009) comparisons. Short‐term increases in carbohydrate availability increased the density of A. gracilipes workers on plants by as much as 400% and reduced tending of honeydew‐excreting insects by this species by up to 89%, with similar responses across years. In contrast, ants at the uninvaded site in 2007 showed a weak and non‐significant forager recruitment response. Across all sites, A. gracilipes workers were the only ants that responded to carbohydrate manipulations in 2009. Furthermore, ant–carbohydrate dynamics at a site newly invaded by A. gracilipes quickly diverged from dynamics at uninvaded sites and converged on those of the site with an established invasion. These findings suggest that carbohydrate resources may be particularly important for A. gracilipes invasions, and underscore the importance of species interactions, particularly putative mutualisms, in facilitating exotic species invasions.     

Is evolution faster at ecotones? A test using rates and tempo of diet transitions in Neotropical Sigmodontinae (Rodentia,Cricetidae)

We evaluated whether evolution is faster at ecotones as niche shifts may be needed to persist under unstable environment. We mapped diet evolution along the evolutionary history of 350 sigmodontine species. Mapping was used in three new tip‐based metrics of trait evolution – Transition Rates, Stasis Time, and Last Transition Time – which were spatialized at the assemblage level (aTR, aST, aTL). Assemblages were obtained by superimposing range maps on points located at core and ecotone of the 93 South American ecoregions. Using Linear Mixed Models, we tested whether ecotones have species with more changes from the ancestral diet (higher aTR), have maintained the current diet for a shorter time (lower aST), and have more recent transitions to the current diet (lower aLT) than cores. We found lower aTR, and higher aST and aLT at ecotones than at cores. Although ecotones are more heterogeneous, both environmentally and in relation to selection pressures they exert on organisms, ecotone species change little from the ancestral diet as generalist habits are necessary toward feeding in ephemeral environments. The need to incorporate phylogenetic uncertainty in tip‐based metrics was evident from large uncertainty detected. Our study integrates ecology and evolution by analyzing how fast trait evolution is across space.     

Can limiting similarity increase invasion resistance? A meta‐analysis of experimental studies

Synthesis We used meta‐analyses to examine experimental evidence that functional similarity between invaders and resident communities reduces invasion. We synthesized evidence from studies that experimentally added seed to resident communities in which the functional group composition had been manipulated. We found communities containing functionally similar resident species reduced invasion of forb but not grass invaders. However, experimental design dramatically influenced the results – with evidence for limiting similarity only found in artificially assembled communities, and not when studies used functional group removal from more ‘natural communities’. We suggest that functional group similarity plays a limited role in biotic resistance in established communities. The principle of limiting similarity suggests that species must be functionally different to coexist; based on the assumption that inter‐specific competition should be greatest between functionally similar species. There has been controversy over the generality of this assembly rule for plant communities with some studies finding evidence for limiting similarity and others not. One approach to testing this is to examine the ‘invasion’ success of species into communities in which the functional group composition has been manipulated. Using a meta‐analysis approach, we examined the generality of limiting similarity for plant communities based on published experimental studies. We asked – is establishment of an invading species less successful if it belongs to a functional group that is already present in the community compared to a community in which that functional group is absent? We explored separately colonisation (i.e. germination, establishment or seedling survival) and performance (i.e. biomass, cover or growth) of different functional groups (forbs and grasses) and experimental designs (removal experiments of more or less natural communities and synthetic‐assemblage experiments). We found that communities containing functionally similar resident species did reduce invader colonisation and performance of forb invaders, but did not reduce colonisation or performance of grass invaders. Evidence in support of limiting similarity was only detected in synthetic‐assemblage experiments and not when studies used functional group removal from ‘natural’ communities. Functional similarity is an important aspect of biotic resistance for forb invaders, but was only found in artificial communities. This has implications for restoration ecology especially when communities are built de novo. However, we suggest that limiting similarity plays a limited role in biotic resistance, because no evidence was detected in established communities.     

Does climate determine broad‐scale patterns of species richness? A test of the causal link by natural experiment

Aim Broad‐scale spatial patterns of species richness are very strongly correlated with climatic variables. If there is a causal link, i.e. if climate directly or indirectly determines patterns of richness, then when the climatic variables change, richness should change in the manner that spatial correlations between richness and climate would predict. The present study tests this prediction using seasonal changes in climatic variables and bird richness. Location We used a grid of equal area quadrats (37 000 km²) covering North and Central America as far south as Nicaragua. Methods Summer and winter bird distribution data were drawn from monographs and field guides. Climatic data came from published sources. We also used remotely sensed NDVI (normalized difference vegetation index — a measure of greenness). Results Bird species richness changes temporally (between summer and winter) in a manner that is close to, but statistically distinguishable from, the change one would predict from models relating the spatial variation in richness at a single time to climatic variables. If one further takes into account the seasonal changes in NDVI and within‐season variability of temperature and precipitation, then winter and summer richness follow congruent, statistically indistinguishable patterns. Main conclusions Our results are consistent with the hypothesis that climatic variables (temperature and precipitation) and vegetation cover directly or indirectly influence patterns of bird species richness.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie

Many clonal plants consist of many connected individual ramets, allowing them to share water and nutrients via physiological integration. Integration among ramets may also improve the ability of clonal plants to tolerate abiotic stress or improve the competitive ability of individual ramets. Here I use a field experiment to determine whether clonal integration improves ramet performance for a widespread clonal tree species invading into native prairie. Aspen (Populus tremuloides) dominates the southern treeline in western Canada, has long-lived belowground connections between mother and daughter ramets, and reproduces vegetatively via resprouting rhizomes after disturbance. I applied two competition treatments (neighbors present or absent) and two clonal integration treatments (belowground rhizomes between mother and daughter ramets either severed or left intact) to 12 replicate Populus daughter ramets at each of three sites. Neighbors improved the survivorship of Populus ramets by 25-35% after 2 yr, but decreased growth by ～20%. Clonal integration tended to improve ramet survival and growth, but these trends were often not significant. Clonal integration did not alter the effects of competition from neighboring vegetation, suggesting that connections between ramets do not necessarily improve the competitive ability of Populus invading into native prairie.     

Did some red alga‐derived plastids evolve via kleptoplastidy? A hypothesis

The evolution of plastids has a complex and still unresolved history. These organelles originated from a cyanobacterium via primary endosymbiosis, resulting in three eukaryotic lineages: glaucophytes, red algae, and green plants. The red and green algal plastids then spread via eukaryote–eukaryote endosymbioses, known as secondary and tertiary symbioses, to numerous heterotrophic protist lineages. The number of these horizontal plastid transfers, especially in the case of red alga‐derived plastids, remains controversial. Some authors argue that the number of plastid origins should be minimal due to perceived difficulties in the transformation of a eukaryotic algal endosymbiont into a multimembrane plastid, but increasingly the available data contradict this argument. I suggest that obstacles in solving this dilemma result from the acceptance of a single evolutionary scenario for the endosymbiont‐to‐plastid transformation formulated by Cavalier‐Smith & Lee (1985). Herein I discuss data that challenge this evolutionary scenario. Moreover, I propose a new model for the origin of multimembrane plastids belonging to the red lineage and apply it to the dinoflagellate peridinin plastid. The new model has several general and practical implications, such as the requirement for a new definition of cell organelles and in the construction of chimeric organisms.     

Does allometry of a sexually selected ornamental trait vary with sexual selection intensity? A multi‐species test in damselflies

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Does size matter? A test of size‐specific mortality in Atlantic salmon Salmo salar smolts tagged with acoustic transmitters

Mortality rates of wild Atlantic salmon Salmo salar smolts implanted with acoustic transmitters were assessed to determine if mortality was size dependent. The routinely accepted, but widely debated, ‘2% transmitter mass: body mass’ rule in biotelemetry was tested by extending the transmitter burden up to 12·7% of body mass in small [mean fork length (L_F) 138·3 mm, range 115–168 mm] downstream migrating S. salar smolts. Over the short timescale of emigration (range 11·9–44·5 days) through the lower river and estuary, mortality was not related to S. salar size, nor was a relationship found between mortality probability and transmitter mass: body mass or transmitter length: L_F ratios. This study provides further evidence that smolt migration studies can deviate from the ‘2% rule’ of thumb, to more appropriate study‐specific measures, which enables the use of fishes representative of the body size in natural populations without undue effects.     

MDA‐7/IL‐24 regulates proliferation,invasion and tumor cell radiosensitivity: A new cancer therapy?

The novel cytokine MDA-7/IL-24 was identified by subtractive hybridization in the mid-1990s as a cytokine whose expression increased during the induction of terminal differentiation, and that was either not expressed or was present at low levels in tumor cells compared to non-transformed cells. Multiple studies from several laboratories have subsequently demonstrated that expression of IL-24 in tumor cells, but not in non-transformed cells, causes their growth arrest and ultimately cell death. In addition, IL-24 has been noted to be a radiosensitizing cytokine, which in part is due to the generation of reactive oxygen species (ROS) and causing endoplasmic reticulum stress. Recent publications of Phase I trial data have shown that a recombinant adenovirus to express MDA-7/IL-24 (Ad.mda-7 (INGN 241)) was safe and had tumoricidal effects in patients, which argues that IL-24 may have therapeutic value. This review describes what is known about the impact of IL-24 on tumor cell biology in addition to approaches that may enhance the toxicity of this novel cytokine.