Antagonistic effects of intraspecific cooperation and interspecific competition on thermal performance.
Tsai Hsiang-Yu,Rubenstein Dustin R,Chen Bo-Fei,Liu Mark,Chan Shih-Fan,Chen De-Pei,Sun Syuan-Jyun,Yuan Tzu-Neng,Shen Sheng-Feng
Understanding how climate-mediated biotic interactions shape thermal niche width is critical in an era of global change. Yet, most previous work on thermal niches has ignored detailed mechanistic information about the relationship between temperature and organismal performance, which can be described by a thermal performance curve. Here, we develop a model that predicts the width of thermal performance curves will be narrower in the presence of interspecific competitors, causing a species' optimal breeding temperature to diverge from that of its competitor. We test this prediction in the Asian burying beetle , confirming that the divergence in actual and optimal breeding temperatures is the result of competition with their primary competitor, blowflies. However, we further show that intraspecific cooperation enables beetles to outcompete blowflies by recovering their optimal breeding temperature. Ultimately, linking abiotic factors and biotic interactions on niche width will be critical for understanding species-specific responses to climate change.
Patterns and drivers of intraspecific variation in avian life history along elevational gradients: a meta-analysis.
Alice Boyle W,Sandercock Brett K,Martin Kathy
Biological reviews of the Cambridge Philosophical Society
Elevational gradients provide powerful natural systems for testing hypotheses regarding the role of environmental variation in the evolution of life-history strategies. Case studies have revealed shifts towards slower life histories in organisms living at high elevations yet no synthetic analyses exist of elevational variation in life-history traits for major vertebrate clades. We examined (i) how life-history traits change with elevation in paired populations of bird species worldwide, and (ii) which biotic and abiotic factors drive elevational shifts in life history. Using three analytical methods, we found that fecundity declined at higher elevations due to smaller clutches and fewer reproductive attempts per year. By contrast, elevational differences in traits associated with parental investment or survival varied among studies. High-elevation populations had shorter and later breeding seasons, but longer developmental periods implying that temporal constraints contribute to reduced fecundity. Analyses of clutch size data, the trait for which we had the largest number of population comparisons, indicated no evidence that phylogenetic history constrained species-level plasticity in trait variation associated with elevational gradients. The magnitude of elevational shifts in life-history traits were largely unrelated to geographic (altitude, latitude), intrinsic (body mass, migratory status), or habitat covariates. Meta-population structure, methodological issues associated with estimating survival, or processes shaping range boundaries could potentially explain the nature of elevational shifts in life-history traits evident in this data set. We identify a new risk factor for montane populations in changing climates: low fecundity will result in lower reproductive potential to recover from perturbations, especially as fewer than half of the species experienced higher survival at higher elevations.
Direct benefits and indirect costs of warm temperatures for high-elevation populations of a solitary bee.
Forrest Jessica R K,Chisholm Sarah P M
Warm temperatures are required for insect flight. Consequently, warming could benefit many high-latitude and high-altitude insects by increasing opportunities for foraging or oviposition. However, warming can also alter species interactions, including interactions with natural enemies, making the net effect of rising temperatures on population growth rate difficult to predict. We investigated the temperature-dependence of nesting activity and lifetime reproductive output over 3 yr in subalpine populations of a pollen-specialist bee, Osmia iridis. Rates of nest provisioning increased with ambient temperatures and with availability of floral resources, as expected. However, warmer conditions did not increase lifetime reproductive output. Lifetime offspring production was best explained by rates of brood parasitism (by the wasp Sapyga), which increased with temperature. Direct observations of bee and parasite activity suggest that although activity of both species is favored by warmer temperatures, bees can be active at lower ambient temperatures, while wasps are active only at higher temperatures. Thus, direct benefits to the bees of warmer temperatures were nullified by indirect costs associated with increased parasite activity. To date, most studies of climate-change effects on pollinators have focused on changing interactions between pollinators and their floral host-plants (i.e., bottom-up processes). Our results suggest that natural enemies (i.e., top-down forces) can play a key role in pollinator population regulation and should not be overlooked in forecasts of pollinator responses to climate change.
Network reorganization and breakdown of an ant-plant protection mutualism with elevation.
Plowman Nichola S,Hood Amelia S C,Moses Jimmy,Redmond Conor,Novotny Vojtech,Klimes Petr,Fayle Tom M
Proceedings. Biological sciences
Both the abiotic environment and the composition of animal and plant communities change with elevation. For mutualistic species, these changes are expected to result in altered partner availability, and shifts in context-dependent benefits for partners. To test these predictions, we assessed the network structure of terrestrial ant-plant mutualists and how the benefits to plants of ant inhabitation changed with elevation in tropical forest in Papua New Guinea. At higher elevations, ant-plants were rarer, species richness of both ants and plants decreased, and the average ant or plant species interacted with fewer partners. However, networks became increasingly connected and less specialized, more than could be accounted for by reductions in ant-plant abundance. On the most common ant-plant, ants recruited less and spent less time attacking a surrogate herbivore at higher elevations, and herbivory damage increased. These changes were driven by turnover of ant species rather than by within-species shifts in protective behaviour. We speculate that reduced partner availability at higher elevations results in less specialized networks, while lower temperatures mean that even for ant-inhabited plants, benefits are reduced. Under increased abiotic stress, mutualistic networks can break down, owing to a combination of lower population sizes, and a reduction in context-dependent mutualistic benefits.
Evolution of plasticity and adaptive responses to climate change along climate gradients.
Kingsolver Joel G,Buckley Lauren B
Proceedings. Biological sciences
The relative contributions of phenotypic plasticity and adaptive evolution to the responses of species to recent and future climate change are poorly understood. We combine recent (1960-2010) climate and phenotypic data with microclimate, heat balance, demographic and evolutionary models to address this issue for a montane butterfly, , along an elevational gradient. Our focal phenotype, wing solar absorptivity, responds plastically to developmental (pupal) temperatures and plays a central role in thermoregulatory adaptation in adults. Here, we show that both the phenotypic and adaptive consequences of plasticity vary with elevation. Seasonal changes in weather generate seasonal variation in phenotypic selection on mean and plasticity of absorptivity, especially at lower elevations. In response to climate change in the past 60 years, our models predict evolutionary declines in mean absorptivity (but little change in plasticity) at high elevations, and evolutionary increases in plasticity (but little change in mean) at low elevation. The importance of plasticity depends on the magnitude of seasonal variation in climate relative to interannual variation. Our results suggest that selection and evolution of both trait means and plasticity can contribute to adaptive response to climate change in this system. They also illustrate how plasticity can facilitate rather than retard adaptive evolutionary responses to directional climate change in seasonal environments.
Flower movement balances pollinator needs and pollen protection.
Haverkamp Alexander,Li Xiang,Hansson Bill S,Baldwin Ian T,Knaden Markus,Yon Felipe
Flower signaling and orientation are key characteristics that determine a flower's pollinator guild. However, many flowers actively move during their daily cycle, changing both their detectability and accessibility to pollinators. The flowers of the wild tobacco Nicotiana attenuata orientate their corolla upward at sunset and downward after sunrise. Here, we investigated the effect of different flower orientations on a major pollinator of N. attenuata, the hawkmoth Manduca sexta. We found that although flower orientation influenced the flight altitude of the moth in respect to the flower, it did not alter the moth's final flower choice. These behavioral observations were consistent with the finding that orientation did not systematically change the spatial distribution of floral volatiles, which are major attractants for the moths. Moreover, hawkmoths invested the same amount of time into probing flowers at different orientations, even though they were only able to feed and gather pollen from horizontally and upward-oriented flowers, but not from downward-facing flowers. The orientation of the flower was hence crucial for a successful interaction between N. attenuata and its hawkmoth pollinator. Additionally, we also investigated potential adverse effects of exposing flowers at different orientations to natural daylight levels, finding that anther temperature of upward-oriented flowers was more than 7°C higher than for downward-oriented flowers. This increase in temperature likely caused the significantly reduced germination success that was observed for pollen grains from upward-oriented flowers in comparison to those of downward and horizontally oriented flowers. These results highlight the importance of flower reorientation to balance pollen protection and a successful interaction of the plant with its insect pollinators by maintaining the association between flower volatiles and flower accessibility to the pollinator.
Linking inter-annual variation in environment, phenology, and abundance for a montane butterfly community.
Stewart James E,Illán Javier Gutiérrez,Richards Shane A,Gutiérrez David,Wilson Robert J
Climate change has caused widespread shifts in species' phenology, but the consequences for population and community dynamics remain unclear because of uncertainty regarding the species-specific drivers of phenology and abundance, and the implications for synchrony among interacting species. Here, we develop a statistical model to quantify inter-annual variation in phenology and abundance over an environmental gradient, and use it to identify potential drivers of phenology and abundance in co-occurring species. We fit the model to counts of 10 butterfly species with single annual generations over a mountain elevation gradient, as an exemplar system in which temporally limited availability of biotic resources and favorable abiotic conditions impose narrow windows of seasonal activity. We estimate parameters describing changes in abundance, and the peak time and duration of the flight period, over ten years (2004-2013) and across twenty sample locations (930-2,050 m) in central Spain. We also use the model outputs to investigate relationships of phenology and abundance with temperature and rainfall. Annual shifts in phenology were remarkably consistent among species, typically showing earlier flight periods during years with warm conditions in March or May-June. In contrast, inter-annual variation in relative abundance was more variable among species, and generally less well associated with climatic conditions. Nevertheless, warmer temperatures in June were associated with increased relative population growth in three species, and five species had increased relative population growth in years with earlier flight periods. These results suggest that broadly coherent interspecific changes to phenology could help to maintain temporal synchrony in community dynamics under climate change, but that the relative composition of communities may vary due to interspecific inconsistency in population dynamic responses to climate change. However, it may still be possible to predict abundance change for species based on a robust understanding of relationships between their population dynamics and phenology, and the environmental drivers of both.
Adaptive strategies of high-flying migratory hoverflies in response to wind currents.
Gao Boya,Wotton Karl R,Hawkes Will L S,Menz Myles H M,Reynolds Don R,Zhai Bao-Ping,Hu Gao,Chapman Jason W
Proceedings. Biological sciences
Large migrating insects, flying at high altitude, often exhibit complex behaviour. They frequently elect to fly on winds with directions quite different from the prevailing direction, and they show a degree of common orientation, both of which facilitate transport in seasonally beneficial directions. Much less is known about the migration behaviour of smaller (10-70 mg) insects. To address this issue, we used radar to examine the high-altitude flight of hoverflies (Diptera: Syrphidae), a group of day-active, medium-sized insects commonly migrating over the UK. We found that autumn migrants, which must move south, did indeed show migration timings and orientation responses that would take them in this direction, despite the unfavourability of the prevailing winds. Evidently, these hoverfly migrants must have a compass (probably a time-compensated solar mechanism), and a means of sensing the wind direction (which may be determined with sufficient accuracy at ground level, before take-off). By contrast, hoverflies arriving in the UK in spring showed weaker orientation tendencies, and did not correct for wind drift away from their seasonally adaptive direction (northwards). However, the spring migrants necessarily come from the south (on warm southerly winds), so we surmise that complex orientation behaviour may not be so crucial for the spring movements.
Bumblebees moving up: shifts in elevation ranges in the Pyrenees over 115 years.
Marshall Leon,Perdijk Floor,Dendoncker Nicolas,Kunin William,Roberts Stuart,Biesmeijer Jacobus C
Proceedings. Biological sciences
In a warming climate, species are expected to shift their geographical ranges to higher elevations and latitudes, and if interacting species shift at different rates, networks may be disrupted. To quantify the effects of ongoing climate change, repeating historical biodiversity surveys is necessary. In this study, we compare the distribution of a plant-pollinator community between two surveys 115 years apart (1889 and 2005-06), reporting distribution patterns and changes observed for bumblebee species and bumblebee-visited plants in the Gavarnie-Gèdre commune in the Pyrenees, located in southwest Europe at the French-Spanish border. The region has warmed significantly over this period, alongside shifts in agricultural land use and forest. The composition of the bumblebee community shows relative stability, but we observed clear shifts to higher elevations for bumblebees (averaging 129 m) and plants (229 m) and provide preliminary evidence that some bumblebee species shift with the plants they visit. We also observe that some species have been able to occupy the same climate range in both periods by shifting elevation range. The results suggest the need for long-term monitoring to determine the role and impact of the different drivers of global change, especially in montane habitats where the impacts of climate changes are anticipated to be more extreme.
Limited tolerance by insects to high temperatures across tropical elevational gradients and the implications of global warming for extinction.
García-Robledo Carlos,Kuprewicz Erin K,Staines Charles L,Erwin Terry L,Kress W John
Proceedings of the National Academy of Sciences of the United States of America
The critical thermal maximum (CTmax), the temperature at which motor control is lost in animals, has the potential to determine if species will tolerate global warming. For insects, tolerance to high temperatures decreases with latitude, suggesting that similar patterns may exist along elevational gradients as well. This study explored how CTmax varies among species and populations of a group of diverse tropical insect herbivores, the rolled-leaf beetles, across both broad and narrow elevational gradients. Data from 6,948 field observations and 8,700 museum specimens were used to map the elevational distributions of rolled-leaf beetles on two mountains in Costa Rica. CTmax was determined for 1,252 individual beetles representing all populations across the gradients. Initial morphological identifications suggested a total of 26 species with populations at different elevations displaying contrasting upper thermal limits. However, compared with morphological identifications, DNA barcodes (cytochrome oxidase I) revealed significant cryptic species diversity. DNA barcodes identified 42 species and haplotypes across 11 species complexes. These 42 species displayed much narrower elevational distributions and values of CTmax than the 26 morphologically defined species. In general, species found at middle elevations and on mountaintops are less tolerant to high temperatures than species restricted to lowland habitats. Species with broad elevational distributions display high CTmax throughout their ranges. We found no significant phylogenetic signal in CTmax, geography, or elevational range. The narrow variance in CTmax values for most rolled-leaf beetles, especially high-elevation species, suggests that the risk of extinction of insects may be substantial under some projected rates of global warming.
Testing for local adaptation and evolutionary potential along altitudinal gradients in rainforest Drosophila: beyond laboratory estimates.
O'Brien Eleanor K,Higgie Megan,Reynolds Alan,Hoffmann Ary A,Bridle Jon R
Global change biology
Predicting how species will respond to the rapid climatic changes predicted this century is an urgent task. Species distribution models (SDMs) use the current relationship between environmental variation and species' abundances to predict the effect of future environmental change on their distributions. However, two common assumptions of SDMs are likely to be violated in many cases: (i) that the relationship of environment with abundance or fitness is constant throughout a species' range and will remain so in future and (ii) that abiotic factors (e.g. temperature, humidity) determine species' distributions. We test these assumptions by relating field abundance of the rainforest fruit fly Drosophila birchii to ecological change across gradients that include its low and high altitudinal limits. We then test how such ecological variation affects the fitness of 35 D. birchii families transplanted in 591 cages to sites along two altitudinal gradients, to determine whether genetic variation in fitness responses could facilitate future adaptation to environmental change. Overall, field abundance was highest at cooler, high-altitude sites, and declined towards warmer, low-altitude sites. By contrast, cage fitness (productivity) increased towards warmer, lower-altitude sites, suggesting that biotic interactions (absent from cages) drive ecological limits at warmer margins. In addition, the relationship between environmental variation and abundance varied significantly among gradients, indicating divergence in ecological niche across the species' range. However, there was no evidence for local adaptation within gradients, despite greater productivity of high-altitude than low-altitude populations when families were reared under laboratory conditions. Families also responded similarly to transplantation along gradients, providing no evidence for fitness trade-offs that would favour local adaptation. These findings highlight the importance of (i) measuring genetic variation in key traits under ecologically relevant conditions, and (ii) considering the effect of biotic interactions when predicting species' responses to environmental change.
Partner abundance controls mutualism stability and the pace of morphological change over geologic time.
Chomicki Guillaume,Renner Susanne S
Proceedings of the National Academy of Sciences of the United States of America
Mutualisms that involve symbioses among specialized partners may be more stable than mutualisms among generalists, and theoretical models predict that in many mutualisms, partners exert reciprocal stabilizing selection on traits directly involved in the interaction. A corollary is that mutualism breakdown should increase morphological rates of evolution. We here use the largest ant-plant clade (Hydnophytinae), with different levels of specialization for mutualistic ant symbionts, to study the ecological context of mutualism breakdown and the response of a key symbiosis-related trait, domatium entrance hole size, which filters symbionts by size. Our analyses support three predictions from mutualism theory. First, all 12 losses apparently only occur from a generalist symbiotic state. Second, mutualism losses occurred where symbionts are scarce, in our system at high altitudes. Third, domatium entrance hole size barely changes in specialized symbiotic species, but evolves rapidly once symbiosis with ants has broken down, with a "morphorate map" revealing that hotspots of entrance hole evolution are clustered in high-altitude areas. Our study reveals that mutualistic strategy profoundly affects the pace of morphological change in traits involved in the interaction and suggests that shifts in partners' relative abundances may frequently drive reversions of generalist mutualisms to autonomy.
Contrasting forms of competition set elevational range limits of species.
Chan Shih-Fan,Shih Wei-Kai,Chang An-Yu,Shen Sheng-Feng,Chen I-Ching
How abiotic and biotic factors constrain distribution limits at the harsh and benign edges of species ranges is hotly debated, partly because macroecological experiments testing the proximate causes of distribution limits are scarce. It has long been recognized - at least since Darwin's On the Origin of Species - that a harsh climate strengthens competition and thus sets species range limits. Using thorough field manipulations along a large elevation gradient, we show the mechanisms by which temperature determines competition type, resulting in a transition from interference to exploitative competition from the lower to the upper elevation limits in burying beetles (Nicrophorus nepalensis). This transition is an example of Darwin's classic hypothesis that benign climates favor direct competition for highly accessible resources while harsh climates result in competition through resources of high rivalry. We propose that identifying the properties of these key resources will provide a more predictive framework to understand the interplay between biotic and abiotic factors in determining geographic range limits.
Life at High Latitudes Does Not Require Circadian Behavioral Rhythmicity under Constant Darkness.
Bertolini Enrico,Schubert Frank K,Zanini Damiano,Sehadová Hana,Helfrich-Förster Charlotte,Menegazzi Pamela
Current biology : CB
Nearly all organisms evolved endogenous self-sustained timekeeping mechanisms to track and anticipate cyclic changes in the environment. Circadian clocks, with a periodicity of about 24 h, allow animals to adapt to day-night cycles. Biological clocks are highly adaptive, but strong behavioral rhythms might be a disadvantage for adaptation to weakly rhythmic environments such as polar areas [1, 2]. Several high-latitude species, including Drosophila species, were found to be highly arrhythmic under constant conditions [3-6]. Furthermore, Drosophila species from subarctic regions can extend evening activity until dusk under long days. These traits depend on the clock network neurochemistry, and we previously proposed that high-latitude Drosophila species evolved specific clock adaptations to colonize polar regions [5, 7, 8]. We broadened our analysis to 3 species of the Chymomyza genus, which diverged circa 5 million years before the Drosophila radiation  and colonized both low and high latitudes [10, 11]. C. costata, pararufithorax, and procnemis, independently of their latitude of origin, possess the clock neuronal network of low-latitude Drosophila species, and their locomotor activity does not track dusk under long photoperiods. Nevertheless, the high-latitude C. costata becomes arrhythmic under constant darkness (DD), whereas the two low-latitude species remain rhythmic. Different mechanisms are behind the arrhythmicity in DD of C. costata and the high-latitude Drosophila ezoana, suggesting that the ability to maintain behavioral rhythms has been lost more than once during drosophilids' evolution and that it might indeed be an evolutionary adaptation for life at high latitudes.
Phenology responses of temperate butterflies to latitude depend on ecological traits.
Faltýnek Fric Zdeněk,Rindoš Michal,Konvička Martin
Global change influences species' seasonal occurrence, or phenology. In cold-adapted insects, the activity is expected to start earlier with a warming climate, but contradictory evidence exists, and the reactions may be linked to species-specific traits. Using data from the GBIF database, we selected 105 single-brooded Holarctic butterflies inhabiting broad latitudinal ranges. We regressed patterns of an adult flight against latitudes of the records, controlling for altitude and year effects. Species with delayed flight periods towards the high latitudes, or stable flight periods across latitudes, prevailed over those that advanced their flight towards the high latitudes. The responses corresponded with the species' seasonality (flight of early season species was delayed and flight of summer species was advanced at high latitudes) and oceanic vs. continental climatic niches (delays in oceanic, stability in continental species). Future restructuring of butterfly seasonal patterns in high latitudes will reflect climatic niches, and hence the evolutionary history of participating species.
Dietary effects on gut microbiota of the mesquite lizard Sceloporus grammicus (Wiegmann, 1828) across different altitudes.
Montoya-Ciriaco Nina,Gómez-Acata Selene,Muñoz-Arenas Ligia Catalina,Dendooven Luc,Estrada-Torres Arturo,Díaz de la Vega-Pérez Aníbal H,Navarro-Noya Yendi E
BACKGROUND:High-altitude ecosystems are extreme environments that generate specific physiological, morphological, and behavioral adaptations in ectotherms. The shifts in gut microbiota of the ectothermic hosts as an adaptation to environmental changes are still largely unknown. We investigated the food ingested and the bacterial, fungal, and protistan communities in feces of the lizard Sceloporus grammicus inhabiting an altitudinal range using metabarcoding approaches. RESULTS:The bacterial phyla Bacteroidetes and Firmicutes, and the genera Bacteroides and Parabacteroides dominated the core fecal bacteriome, while Zygomycota and Ascomycota, and the species Basidiobolus ranarum and Basidiobolus magnus dominated the core fecal mycobiome. The diet of S. grammicus included 29 invertebrate families belonging to Arachnida, Chilopoda, and Insecta. The diversity and abundance of its diet decreased sharply at high altitudes, while the abundance of plant material and Agaricomycetes was significantly higher at the highest site. The composition of the fecal microbiota of S. grammicus was different at the three altitudes, but not between females and males. Dietary restriction in S. grammicus at 4150 m might explain the high fecal abundance of Akkermansia and Oscillopira, bacteria characteristic of long fasting periods, while low temperature favored B. magnus. A high proportion of bacterial functions were digestive in S. grammicus at 2600 and 3100, while metabolism of aminoacids, vitamins, and key intermediates of metabolic pathways were higher at 4150 m. Different assemblages of fungal species in the lizard reflect differences in the environments at different elevations. Pathogens were more prevalent at high elevations than at the low ones. CONCLUSIONS:Limiting food resources at high elevations might oblige S. grammicus to exploit other food resources and its intestinal microbiota have degradative and detoxifying capacities. Sceloporus grammicus might have acquired B. ranarum from the insects infected by the fungus, but its commensal relationship might be established by the quitinolytic capacities of B. ranarum. The mycobiome participate mainly in digestive and degradative functions while the bacteriome in digestive and metabolic functions.
Genomics Reveals Widespread Ecological Speciation in Flightless Insects.
McCulloch Graham A,Foster Brodie J,Dutoit Ludovic,Harrop Thomas W R,Guhlin Joseph,Dearden Peter K,Waters Jonathan M
Recent genomic analyses have highlighted parallel divergence in response to ecological gradients, but the extent to which altitude can underpin such repeated speciation remains unclear. Wing reduction and flight loss have apparently evolved repeatedly in montane insect assemblages, and have been suggested as important drivers of hexapod diversification. We test this hypothesis using genomic analyses of a widespread wing-polymorphic stonefly species complex in New Zealand. We identified over 50,000 polymorphic genetic markers generated across almost 200 Zelandoperla fenestrata stonefly specimens using a newly generated plecopteran reference genome, to reveal widespread parallel speciation between sympatric full-winged and wing-reduced ecotypes. Rather than the existence of a single, widespread, flightless taxon (Zelandoperla pennulata), evolutionary genomic data reveal that wing-reduced upland lineages have speciated repeatedly and independently from full-winged Z. fenestrata. This repeated evolution of reproductive isolation between local ecotype pairs that lack mitochondrial DNA differentiation suggests that ecological speciation has evolved recently. A cluster of outlier SNPs detected in independently wing-reduced lineages, tightly linked in an approximately 85 kb genomic region that includes the developmental 'supergene' doublesex, suggests that this 'island of divergence' may play a key role in rapid ecological speciation.
Ecology of insect host-parasitoid communities.
Force D C
Science (New York, N.Y.)
Although conclusive evidence is lacking for its establishment, the thesis that complexity adds stability to communities is probably accepted by the majority of ecologists. I believe this attitude found its origins in the indisputable fact that there are latitudinal and altitudinal changes in community complexity. As one progresses northward or southward from the equator, or higher in altitude in most parts of the world, one cannot help but notice that communities tend to become simpler, that is, there are fewer species per community. At the same time, these communities appear to become less stable. But perhaps this change in stability is in appearance only; they appear to be less stable because of the relatively greater number of individuals comprising each species population in temperate areas. Each population, because of its greater numbers, is therefore conspicuous, and changes in these numbers are noticed. We are particularly aware of such changes because populations in these areas of the world have been comparatively well studied. Many of the most studied populations include species of economic importance where changes in population numbers are vital to agricultural or forestry practices. Equatorial populations, on the other hand, contain smaller numbers of individuals of each species because of the greater number of species present. Number changes are simply not as noticeable because the population itself is not as obvious among the other populations. It may be that when (if ever) we have as much data on equatorial populations as we have on those of temperate climates, we will find fluctuations of equal relative magnitude (but not of equal numbers, of course). If, on the other hand, we really do find a correlation between complexity and stability, the suggestion by May (12) that stability permits complexity may be well worth investigating. Because of its organization and physical setting, the Rhopalomyia community I have studied might be expected to have considerable stability. In fact, however, it does not. Each of the populations in the community fluctuates greatly and irregularly in both percentages and numbers, and these populations apparently become locally extinct occasionally, because they sometimes cannot be found even in extensive collections. After studying several of the more important parasitoid species, it is evident to me that there is little or nothing about their interactions that might induce greater community stability. Each species seems to have evolved into the community with no higher purpose than simply to usurp what it can from some other member, and it does this by concentrating its energies on better competitive mechanisms rather than higher reproductive capacities. There are never empty niches to be filled by organisms having the "correct specifications" because new niches are created out of parts of older, broader niches which were occupied by other, more r-selected organisms. Thus, perhaps we have read too much into community organization. Perhaps the "ifiling of niches" is essentially nothing more than the haphazard result of competitive jostling among species; and that as communities develop, they are not necessarily programmed for such things as greater stability or better energy utilizationthe species merely become more closely packed.
Altitudinal changes in malaria incidence in highlands of Ethiopia and Colombia.
Siraj A S,Santos-Vega M,Bouma M J,Yadeta D,Ruiz Carrascal D,Pascual M
Science (New York, N.Y.)
The impact of global warming on insect-borne diseases and on highland malaria in particular remains controversial. Temperature is known to influence transmission intensity through its effects on the population growth of the mosquito vector and on pathogen development within the vector. Spatiotemporal data at a regional scale in highlands of Colombia and Ethiopia supplied an opportunity to examine how the spatial distribution of the disease changes with the interannual variability of temperature. We provide evidence for an increase in the altitude of malaria distribution in warmer years, which implies that climate change will, without mitigation, result in an increase of the malaria burden in the densely populated highlands of Africa and South America.
Shared Genetic Signals of Hypoxia Adaptation in Drosophila and in High-Altitude Human Populations.
Jha Aashish R,Zhou Dan,Brown Christopher D,Kreitman Martin,Haddad Gabriel G,White Kevin P
Molecular biology and evolution
The ability to withstand low oxygen (hypoxia tolerance) is a polygenic and mechanistically conserved trait that has important implications for both human health and evolution. However, little is known about the diversity of genetic mechanisms involved in hypoxia adaptation in evolving populations. We used experimental evolution and whole-genome sequencing in Drosophila melanogaster to investigate the role of natural variation in adaptation to hypoxia. Using a generalized linear mixed model we identified significant allele frequency differences between three independently evolved hypoxia-tolerant populations and normoxic control populations for approximately 3,800 single nucleotide polymorphisms. Around 50% of these variants are clustered in 66 distinct genomic regions. These regions contain genes that are differentially expressed between hypoxia-tolerant and normoxic populations and several of the differentially expressed genes are associated with metabolic processes. Additional genes associated with respiratory and open tracheal system development also show evidence of directional selection. RNAi-mediated knockdown of several candidate genes' expression significantly enhanced survival in severe hypoxia. Using genomewide single nucleotide polymorphism data from four high-altitude human populations-Sherpas, Tibetans, Ethiopians, and Andeans, we found that several human orthologs of the genes under selection in flies are also likely under positive selection in all four high-altitude human populations. Thus, our results indicate that selection for hypoxia tolerance can act on standing genetic variation in similar genes and pathways present in organisms diverged by hundreds of millions of years.
Humboldt's enigma: What causes global patterns of mountain biodiversity?
Rahbek Carsten,Borregaard Michael K,Colwell Robert K,Dalsgaard Bo,Holt Ben G,Morueta-Holme Naia,Nogues-Bravo David,Whittaker Robert J,Fjeldså Jon
Science (New York, N.Y.)
Mountains contribute disproportionately to the terrestrial biodiversity of Earth, especially in the tropics, where they host hotspots of extraordinary and puzzling richness. With about 25% of all land area, mountain regions are home to more than 85% of the world's species of amphibians, birds, and mammals, many entirely restricted to mountains. Biodiversity varies markedly among these regions. Together with the extreme species richness of some tropical mountains, this variation has proven challenging to explain under traditional climatic hypotheses. However, the complex climatic characteristics of rugged mountain regions differ fundamentally from those of lowland regions, likely playing a key role in generating and maintaining diversity. With ongoing global changes in climate and land use, the role of mountains as refugia for biodiversity may well come under threat.
Windborne long-distance migration of malaria mosquitoes in the Sahel.
Huestis Diana L,Dao Adama,Diallo Moussa,Sanogo Zana L,Samake Djibril,Yaro Alpha S,Ousman Yossi,Linton Yvonne-Marie,Krishna Asha,Veru Laura,Krajacich Benjamin J,Faiman Roy,Florio Jenna,Chapman Jason W,Reynolds Don R,Weetman David,Mitchell Reed,Donnelly Martin J,Talamas Elijah,Chamorro Lourdes,Strobach Ehud,Lehmann Tovi
Over the past two decades efforts to control malaria have halved the number of cases globally, yet burdens remain high in much of Africa and the elimination of malaria has not been achieved even in areas where extreme reductions have been sustained, such as South Africa. Studies seeking to understand the paradoxical persistence of malaria in areas in which surface water is absent for 3-8 months of the year have suggested that some species of Anopheles mosquito use long-distance migration. Here we confirm this hypothesis through aerial sampling of mosquitoes at 40-290 m above ground level and provide-to our knowledge-the first evidence of windborne migration of African malaria vectors, and consequently of the pathogens that they transmit. Ten species, including the primary malaria vector Anopheles coluzzii, were identified among 235 anopheline mosquitoes that were captured during 617 nocturnal aerial collections in the Sahel of Mali. Notably, females accounted for more than 80% of all of the mosquitoes that we collected. Of these, 90% had taken a blood meal before their migration, which implies that pathogens are probably transported over long distances by migrating females. The likelihood of capturing Anopheles species increased with altitude (the height of the sampling panel above ground level) and during the wet seasons, but variation between years and localities was minimal. Simulated trajectories of mosquito flights indicated that there would be mean nightly displacements of up to 300 km for 9-h flight durations. Annually, the estimated numbers of mosquitoes at altitude that cross a 100-km line perpendicular to the prevailing wind direction included 81,000 Anopheles gambiae sensu stricto, 6 million A. coluzzii and 44 million Anopheles squamosus. These results provide compelling evidence that millions of malaria vectors that have previously fed on blood frequently migrate over hundreds of kilometres, and thus almost certainly spread malaria over these distances. The successful elimination of malaria may therefore depend on whether the sources of migrant vectors can be identified and controlled.
Artificial intelligence reveals environmental constraints on colour diversity in insects.
Wu Shipher,Chang Chun-Min,Mai Guan-Shuo,Rubenstein Dustin R,Yang Chen-Ming,Huang Yu-Ting,Lin Hsu-Hong,Shih Li-Cheng,Chen Sheng-Wei,Shen Sheng-Feng
Explaining colour variation among animals at broad geographic scales remains challenging. Here we demonstrate how deep learning-a form of artificial intelligence-can reveal subtle but robust patterns of colour feature variation along an ecological gradient, as well as help identify the underlying mechanisms generating this biogeographic pattern. Using over 20,000 images with precise GPS locality information belonging to nearly 2,000 moth species from Taiwan, our deep learning model generates a 2048-dimension feature vector that accurately predicts each species' mean elevation based on colour and shape features. Using this multidimensional feature vector, we find that within-assemblage image feature variation is smaller in high elevation assemblages. Structural equation modeling suggests that this reduced image feature diversity is likely the result of colder environments selecting for darker colouration, which limits the colour diversity of assemblages at high elevations. Ultimately, with the help of deep learning, we will be able to explore the endless forms of natural morphological variation at unpreceded depths.