Age‐dependent mutational effects curtail the evolution of senescence by antagonistic pleiotropy

One of the two main hypotheses to account for ageing is antagonistic pleiotropy (AP). This model requires alleles that increase vital rates (reproduction or survival) at early age at the expense of vital rates at late age. An important focus of evolutionary studies has been to assess the relative abundance of AP‐type aging alleles that arise through mutation. Here, we develop theory that predicts that senescence per se reduces the probability that these alleles arise by mutation. A direct result is that these mutations should arise with extremely low frequencies in already senescing populations. This has profound implications for the evolution of life histories because it implies that the adaptive evolution of aging via AP will experience negative feedback. This theory also clarifies the previously inexplicable epistatic patterns of genetic covariance across age‐specific vital rates that are observed in mutation accumulation experiments. We show that this epistasis is an emergent property of aging.     

Infection susceptibility and immune senescence with advancing age replicated in accelerated aging LmnaDhe mice

Aging confers increased susceptibility to common pathogens including influenza A virus. Despite shared vulnerability to infection with advancing age in humans and rodents, the relatively long time required for immune senescence to take hold practically restricts the use of naturally aged mice to investigate aging‐induced immunological shifts. Here, we show accelerated aging Lmna^Dhe mice with spontaneous mutation in the nuclear scaffolding protein, lamin A, replicate infection susceptibility, and substantial immune cell shifts that occur with advancing age. Naturally aged (≥20 month) and 2‐ to 3‐month‐old Lmna^Dhe mice share near identically increased influenza A susceptibility compared with age‐matched Lmna^WT control mice. Increased mortality and higher viral burden after influenza infection in Lmna^Dhe mice parallel reduced accumulation of lung alveolar macrophage cells, systemic expansion of immune suppressive Foxp3⁺ regulatory T cells, and skewed immune dominance among viral‐specific CD8⁺ T cells similar to the immunological phenotype of naturally aged mice. Thus, aging‐induced infection susceptibility and immune senescence are replicated in accelerated aging Lmna^Dhe mice.     

Age at sexual maturity of male and female banana weevils,Cosmopolites sordidus Germar (Coleoptera: Curculionidae)

This study was to determine the age at sexual maturity and the relationships between age and internal reproductive organs of Cosmopolites sordidus. Male banana weevils become sexually mature 18 days after emergence (DAE), that is after 2 weeks of adult eclosion, in spite of the fact that spermatogenesis is completed at emergence. A positive correlation exists between age and male internal organs, for example for mean testis diameter (r = .849, p ≤ .001), mean seminal vesicle diameter (r = .679, p ≤ .001), mean accessory gland length (r = .561, p ≤ .01) and mean accessory gland diameter (r = .498, p ≤ .05), respectively. Significant differences were recorded between sexually mature and immature weevils with respect to mean testis diameter (T = 4.471, p ≤ .001) and mean seminal vesicle diameter (T = 3.939, p ≤ .001), but not with mean accessory gland length and mean accessory gland diameter (T = 1.899 and 1.661). Male internal organs were visibly underdeveloped at emergence but became significantly enlarged and developed on attainment of sexual maturity. Female C. sordidus, on the other hand, are sexually mature at 5 DAE. There was also a strong, positive correlation between age of females and mean ovariole length (r = .656, p ≤ .001), and significant differences existed between mean ovariole lengths of sexually mature and immature females (T = 4.306, p ≤ .001). Increasing age of females witnessed progressive increases in ovariole lengths and developmental changes within female ovarioles and calyces. The findings made here may be helpful in Musa germplasm screening works, as weevils bred on susceptible cultivars may reveal similar results, while those bred on resistant ones may experience possible delays in their reproductive developments.     

Allelopathic effects of Alexandrium fundyense (Dinophyceae) on Thalassiosira cf. gravida (Bacillariophyceae): a matter of size

Allelopathic interactions among phytoplankton are well documented. The potency of allelopathic species and responses of target species to allelochemicals are quite variable, however, limiting full understanding of the role these interactions may play in nature. One trait that may influence the sensitivity of an individual to allelochemicals is cell size. The few studies that have examined relationships between cell size and susceptibility to allelochemicals have compared different species and thus could not distinguish between the role of size and species‐specific physiological differences. Culturing an actively sexually reproducing diatom allowed us to focus on the influence of target cell size within a single species. We studied growth and nutrient acquisition by the chain‐forming Thalassiosira cf. gravida Clever in the presence and absence of allelochemicals released by Alexandrium fundyense Balech as a function of T. cf. gravida cell size. Upon exposure to filtrate of A. fundyense, T. cf. gravida cultures “bleached” and both growth and nutrient utilization ceased for up to 4 d. The magnitude of the effect was dependent on filtrate concentration and T. cf. gravida cell surface area:volume ratio. The greatest inhibition was observed on the smallest cells, while T. cf. gravida cultures that had undergone cell enlargement via sexual reproduction were least sensitive to A. fundyense filtrate. These results demonstrate that competitor cell size, independent from taxonomy, may influence the outcome of allelopathic interactions. The findings presented here suggest a potential ecological impact of diatom cell size reduction and sexual reproduction that has not yet been described and that may be important in determining diatom survival and success.     

Clonal expansion of mitochondrial DNA deletions is a private mechanism of aging in long‐lived animals

Disruption of mitochondrial metabolism and loss of mitochondrial DNA (mtDNA) integrity are widely considered as evolutionarily conserved (public) mechanisms of aging (López‐Otín et al., Cell, 153, 2013 and 1194). Human aging is associated with loss in skeletal muscle mass and function (Sarcopenia), contributing significantly to morbidity and mortality. Muscle aging is associated with loss of mtDNA integrity. In humans, clonally expanded mtDNA deletions colocalize with sites of fiber breakage and atrophy in skeletal muscle. mtDNA deletions may therefore play an important, possibly causal role in sarcopenia. The nematode Caenorhabditis elegans also exhibits age‐dependent decline in mitochondrial function and a form of sarcopenia. However, it is unclear if mtDNA deletions play a role in C. elegans aging. Here, we report identification of 266 novel mtDNA deletions in aging nematodes. Analysis of the mtDNA mutation spectrum and quantification of mutation burden indicates that (a) mtDNA deletions in nematode are extremely rare, (b) there is no significant age‐dependent increase in mtDNA deletions, and (c) there is little evidence for clonal expansion driving mtDNA deletion dynamics. Thus, mtDNA deletions are unlikely to drive the age‐dependent functional decline commonly observed in C. elegans. Computational modeling of mtDNA dynamics in C. elegans indicates that the lifespan of short‐lived animals such as C. elegans is likely too short to allow for significant clonal expansion of mtDNA deletions. Together, these findings suggest that clonal expansion of mtDNA deletions is likely a private mechanism of aging predominantly relevant in long‐lived animals such as humans and rhesus monkey and possibly in rodents.     

Melanocortin 1 receptor (MC1R) polymorphisms’ influence on size and dermoscopic features of nevi

The melanocortin 1 receptor (MC1R) is a highly polymorphic gene. The loss‐of‐function MC1R variants (“R”) have been strongly associated with red hair color phenotype and an increased melanoma risk. We sequenced the MC1R gene in 175 healthy individuals to assess the influence of MC1R on nevus phenotype. We identified that MC1R variant carriers had larger nevi both on the back [p‐value = .016, adjusted for multiple parameters (adj. p‐value)] and on the upper limbs (adj. p‐value = .007). Specifically, we identified a positive association between the “R” MC1R variants and visible vessels in nevi [p‐value = .033, corrected using the FDR method for multiple comparisons (corrected p‐value)], dots and globules in nevi (corrected p‐value = .033), nevi with eccentric hyperpigmentation (corrected p‐value = .033), a high degree of freckling (adj. p‐value = .019), and an associative trend with presence of blue nevi (corrected p‐value = .120). In conclusion, the MC1R gene appears to influence the nevus phenotype.     

Leaf optical properties during and after drought stress in triticale and maize genotypes differing in drought tolerance

The effect of a short (7 days) and prolonged (14 days) soil drought (D) on leaf optical properties (R reflectance, T transmittance and A absorbance) in PAR and NIR range of irradiation, and on changes in leaf water potential (ψ), leaf injury index (LI), leaf thickness (LT) and chlorophyll (a + b) content (Chl) was studied for maize and triticale genotypes differing in drought tolerance. Under control conditions (C) leaves of maize in comparison to triticale were better hydrated, were thicker and had higher content of chlorophyll (a + b). In non-stressed plants, small differences were observed in measurements of R, T and A. In the range from 500 to 600 nm, the differences between D-resistant and D-sensitive were observed only in transmittance (T) and in range from 700 to 1,100 nm in absorbance (A). In genotypes belonging to the group of D-sensitive T in PAR range and A in NIR range were two times higher than in D-resistant ones. However, in NIR range R for D-sensitive genotypes was lower than for D-resistant ones. The drought stress caused the decrease in ψ, Chl, LT and the increase in leaf injury index (LI). Soil drought applied within 14 days caused larger changes in these physiological characters in comparison to 7 days drought. The observed harmful influence of drought was more visible for maize than triticale. Moreover for genotypes belonging to D-sensitive ones, changes were larger than for D-resistant ones. Similar to changes in ψ, LT and LI drought stress caused changes in leaf optical properties parameters R, A and T. In the PAR range, the highest changes were observed in R, whereas changes in T and A, which were not considerable. Both in maize and triticale, increase in R was higher in plants subjected to 14 days drought than in plants exposed to drought for 7 days. In maize, increase in R was larger for D-sensitive genotype. For both species, changes in T and A of PAR range were small. In NIR range, an increase in R and A, and decrease in T were observed. After 7 days of recovery in plants subjected to shorter period of drought significant differences were still visible in most cases. The same was observed for ψ, LT, LI and Chl parameters. It shows that the period of 7 days rehydration is too short to remove the injuries caused by drought stress. This results indicate that measurements of R, T and A might be useful in practical application for the estimation of the drought tolerance level. Some limitations in the practical application for plant breeding may be caused by relatively high cost of necessary equipment.     

Effect of pollen provision on life-history parameters of phytoseiid predators under hot and dry environmental conditions

Biological control can be severely disrupted under climate change conditions. This is the case of the spider mite Tetranychus urticae in Spanish citrus orchards, where the omnivorous phytoseiid Euseius stipulatus, the most abundant predator in the system, was highly impacted by hot and dry conditions mimicking future warmer summers. Such a situation can often be compensated by the provision of alternative food to support generalist predators. As a first step to studying whether such a technique could be applied in this case, we studied at laboratory conditions whether pollen could mitigate the negative effects of hotter and drier conditions derived of climate change on three phytoseiids with different diet specializations. In addition to E. stipulatus, these predators, which all together, are considered key for the biological control of T. urticae in citrus, are Neoseiulus californicus and Phytoseiulus persimilis. Our results confirm the extremely fine-tuning of T. urticae to hot–dry conditions. They also provide evidence of the poor performance of E. stipulatus, especially in terms of reproduction, compared to the other two phytoseiids at these conditions, even when high-quality pollen was available. Moreover, access to pollen in combination with T. urticae eggs enhanced survival but reduced predation and oviposition relative to a T. urticae-only diet for N. californicus and P. persimilis. Therefore, whether the overall effect of pollen would justify its use in citrus to counteract the deleterious effects of a hotter and drier climate on the natural regulation of T. urticae is still controversial.     

Senescence in growth and reproductive allocation in a bunchgrass

• Senescence is a puzzling phenomenon. Few convincing studies of senescence in perennial herbaceous plants exist. While ramets are known to senesce, whether senescence of bunchgrasses actually occurs is not clear.
• In this study, we grew a set of plants of Elymus excelsus, a bunchgrass, to examine plant size, sexual reproduction and bud formation in individual plants in relation to their gradual ageing, in order to determine whether E. excelsus experiences senescence. We collected data in two consecutive years (2009 and 2010) from field samples of plants from 1 to 5 years old. Using regression models, we performed age‐related analyses of growth and reproduction parameters.
• Our results showed that individual plant size (diameter, individual biomass), total biomass of ramets, number and biomass of reproductive ramets, percentage of ramets that were reproductive, reproductive allocation, over‐wintering buds and juvenile ramets all declined with age. However, vegetative growth (number and biomass of vegetative ramets) did not decrease with age.
• Those plants that survived, dwindled in size as they aged. However, no plants shifted their resource allocation between growth and reproduction as they aged, so the shift in allocation did not account for the fall in size.

     

Antifungal activity of essential oils against three vegetative-compatibility groups of <Emphasis Type="Italic">Verticillium dahliae</Emphasis>

The antifungal activities of volatile phase effects of essential oils from Origanum onites, O. syriacum, O. minutiflorum, O. vulgare, O, marjorana, Thymus vulgaris, T. serpyllum, Rosmarinus officinalis, Salvia officinalis and Micromeria fruticosa were evaluated for their ability to inhibit growth of three vegetative compatibility groups (VCGs) of Verticillium dahliae. Carvacrol was the main component of O. onites, O. minutiflorum and O. vulgare essential oils, while γ-terpinene was the main component of O. syriacum. P-cymene and thymol were the dominant component of T. vulgaris and T. serpyllum. β- thujone and l-camphor were the main component of S. officinalis. Polegone and isomenthone were the dominant components of M. fruticosa essential oil. Based on the in vitro test, the degree of fungistatical effects can be ranked in the following order of inhibition: O. syriacum = O. onites = O. minutiflorum = O. vulgare = T. vulgaris > T. serpyllum > M. fruticosa > S. officinalis = O. marjorana > R. officinalis. The essential oils of S. officinalis, O. marjorana and R. officinalis displayed moderate antifungal activity, that increased with increasing concentrations. Among the VCGs, VCG2A and VCG4B were found to be highly sensitive to the essential oils. The essential oils of O. syriacum, O. onites, O. minutiflorum, O. vulgare and T. vulgaris were the most efficacious, demonstrating strong antifungal activity against all of the tested VCGs of V. dahliae at relatively low concentrations and they could find practical application as natural fungicides in the prevention and protection of plants from V. dahliae infections.     

Hemoglobin senses body temperature

When aspirating human red blood cells (RBCs) into 1.3 μm pipettes (ΔP = −2.3 kPa), a transition from blocking the pipette below a critical temperature T _c = 36.3 ± 0.3°C to passing it above the T _c occurred (micropipette passage transition). With a 1.1 μm pipette no passage was seen which enabled RBC volume measurements also above T _c. With increasing temperature RBCs lost volume significantly faster below than above a T _c = 36.4 ± 0.7 (volume transition). Colloid osmotic pressure (COP) measurements of RBCs in autologous plasma (25°C ≤ T ≤ 39.5°C) showed a T _c at 37.1 ± 0.2°C above which the COP rapidly decreased (COP transition). In NMR T₁-relaxation time measurements, the T₁ of RBCs in autologous plasma changed from a linear (r = 0.99) increment below T _c = 37 ± 1°C at a rate of 0.023 s/K into zero slope above T _c (RBC T₁ transition). In conclusion: An amorphous hemoglobin–water gel formed in the spherical trail, the residual partial sphere of the aspirated RBC. At T _c, a sudden fluidization of the gel occurs. All changes mentioned above happen at a distinct T _c close to body temperature. The T _c is moved +0.8°C to higher temperatures when a D₂O buffer is used. We suggest a mechanism similar to a “glass transition” or a “colloidal phase transition”. At T _c, the stabilizing Hb bound water molecules reach a threshold number enabling a partial Hb unfolding. Thus, Hb senses body temperature which must be inscribed in the primary structure of hemoglobin and possibly other proteins. This article is dedicated to Ludwig Artmann who died on July 21, 2001 on a beautiful summer day during which we performed experiments far away. Ludwig Artmann was a man who encouraged us to be strong and to study hard no matter what were the costs.     

Phosphorus retention as a function of external loading,hydraulic turnover time,area and relative depth in 54 lakes and reservoirs

We analysed phosphorus retention as a function of external loading, hydraulic turnover time, area and relative depth on the basis of published data from 54 lakes and reservoirs in different climate regions around the world. Our analysis demonstrated that reservoirs and lakes that received higher areal loading of phosphorus (TP_in) also retained more P per m² but the proportion of the external P loading retained in the waterbody (retention coefficient, R _P) remained generally independent of TP_in. The waterbodies with longer hydraulic residence times (T _R) retained larger proportions of external P and the correlation between R _P and T _R was much stronger in lakes with areas larger than 25 km² than in the whole data set. TP_in and T _R together determined 78% of the variation in R _P in large lakes. We also partially confirmed our hypothesis that waterbodies with bigger relative depths (Z _R) retain more of the external phosphorus than larger and shallower waterbodies with lower Z _R. The hypothesis was, however, validated only for lakes larger than 25 km² and for those with T _R <0.3 year, where R _P increased significantly with increasing Z _R. In stratified lakes, increasing relative depth correlated with reduced P retention capacity, demonstrating the complex nature of phosphorus biogeochemistry in lake ecosystems.     

Growth characteristics of Uranoscopus scaber Linnaeus 1758, inhabiting the Iskenderun Bay (Northeastern Mediterranean)

This study aimed to determine the age and some growth characteristics of Atlantic stargazer (Uranoscopus scaber) from Iskenderun Bay (Northeastern Mediterranean). For this purpose, a total of 150 Atlantic stargazer ranging in size from 9.1 to 28.0 cm in total length (weight: 11.7–345.7 g) were collected as by-catch from a commercial trawl fishing boat at a depth of 80–100 m between May 2015 and January 2016. The bottom trawl gear used was equipped with a 44 mm stretched mesh size net at the cod-end. The percentage of females and males were 46.7% and 53.3% respectively. The total length–weight relationships equation with coefficient of determination (R²) were found as W = 0.011*TL^3.131, R² = .9728, for all individuals, W = 0.015*TL^3.021, R² = .9512 for females and W = 0.0102*TL^3.136, R² = .9553 for males. By using the von Bertalanffy equation, the growth parameters of Atlantic stargazer were estimated to be L_∞ = 42.35 cm, k = 0.098, t₀ = −1.8474 for all individuals; L_∞ = 36.92 cm, k = 0.138, t₀ = −1.2693 for females and L_∞ = 38.77 cm, k = 0.1, t₀ = −2.334 for males. In this study, age reading was done by two independent readers and index of average percentage error (IAPE) was calculated as 6.1%. The highest condition factor value calculated as 1.81 in the age group 6 and the lowest condition factor value was calculated as 1.48 in the age group 1.     

Thermal acclimation of shoot respiration in an Arctic woody plant species subjected to 22 years of warming and altered nutrient supply

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long‐term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short‐term temperature (T) response and sensitivity of leaf respiration (R), the high‐T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q₁₀); overall, Q₁₀ values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high‐resolution measurements of R across a range of measuring Ts (25–70 °C) yielded insights into the T at which maximal rates of R occurred (T_max). Although growth temperature did not affect T_max, N+P fertilization increased T_max values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.     

Root volatiles in plant–plant interactions II: Root volatiles alter root chemistry and plant–herbivore interactions of neighbouring plants

Volatile organic compounds (VOCs) emitted by plant roots can influence the germination and growth of neighbouring plants. However, little is known about the effects of root VOCs on plant–herbivore interactions of neighbouring plants. The spotted knapweed (Centaurea stoebe) constitutively releases high amounts of sesquiterpenes into the rhizosphere. Here, we examine the impact of C. stoebe root VOCs on the primary and secondary metabolites of sympatric Taraxacum officinale plants and the resulting plant‐mediated effects on a generalist root herbivore, the white grub Melolontha melolontha. We show that exposure of T. officinale to C.stoebe root VOCs does not affect the accumulation of defensive secondary metabolites but modulates carbohydrate and total protein levels in T. officinale roots. Furthermore, VOC exposure increases M. melolontha growth on T. officinale plants. Exposure of T. officinale to a major C. stoebe root VOC, the sesquiterpene (E)‐β‐caryophyllene, partially mimics the effect of the full root VOC blend on M. melolontha growth. Thus, releasing root VOCs can modify plant–herbivore interactions of neighbouring plants. The release of VOCs to increase the susceptibility of other plants may be a form of plant offense.     

SPIKEPIPE: A metagenomic pipeline for the accurate quantification of eukaryotic species occurrences and intraspecific abundance change using DNA barcodes or mitogenomes

The accurate quantification of eukaryotic species abundances from bulk samples remains a key challenge for community ecology and environmental biomonitoring. We resolve this challenge by combining shotgun sequencing, mapping to reference DNA barcodes or to mitogenomes, and three correction factors: (a) a percent‐coverage threshold to filter out false positives, (b) an internal‐standard DNA spike‐in to correct for stochasticity during sequencing, and (c) technical replicates to correct for stochasticity across sequencing runs. The SPIKEPIPE pipeline achieves a strikingly high accuracy of intraspecific abundance estimates (in terms of DNA mass) from samples of known composition (mapping to barcodes R² = .93, mitogenomes R² = .95) and a high repeatability across environmental‐sample replicates (barcodes R² = .94, mitogenomes R² = .93). As proof of concept, we sequence arthropod samples from the High Arctic, systematically collected over 17 years, detecting changes in species richness, species‐specific abundances, and phenology. SPIKEPIPE provides cost‐efficient and reliable quantification of eukaryotic communities.     

Identifying non-invasible habitats for marine copepods using temperature-dependent <Emphasis Type="Italic">R</Emphasis><Subscript>0</Subscript>

If a non-indigenous species is to thrive and become invasive it must first persist under its new set of environmental conditions. Net reproductive rate (R ₀) represents the average number of female offspring produced by a female over its lifetime, and has been used as a metric of population persistence. We modeled R ₀ as a function of ambient water temperature (T) for the invasive marine calanoid copepod Pseudodiaptomus marinus, which is introduced to west coast of North America from East Asia by ship ballast water. The model was based on temperature-dependent stage-structured population dynamics given by a system of ordinary differential equations. We proposed a methodology to identify habitats that are non-invasible for P. marinus using the threshold of R ₀(T) < 1 in order to identify potentially invasible habitats. We parameterized the model using published data on P. marinus and applied R ₀(T) to identify the range of non-invasible habitats in a global scale based on sea surface temperature data. The model predictions matched the field evidence of species occurrences well.     

Quantification of the Effect of Environmental Factors on Seed Germination and Seedling Growth of Eruca (Eruca sativa) Using Mathematical Models

Eruca (Eruca sativa; Brassicaceae) is an important industrial crop due to its ability to grow under a wide range of climatic conditions and in poor fertility lands and also for the quality of seed oil and protein. Seed germination (SG) is an important event in plant’s life history which can significantly be influenced by several environmental factors such as temperature (T), water potential (ψ), salinity, pH, and burial depth. Therefore, this study aimed (i) to investigate the effects of these environmental factors on SG behavior of Eruca using several mathematical models, (ii) to determine the cardinal Ts and tolerance threshold value for each trait (i.e., 50% reduction than its maximum value) affected by the environmental factor, and (iii) to quantify the response of Eruca seedling growth to each environmental factor. The results indicated that Eruca SG and seedling growth were significantly influenced by these factors (P < 0.05). The estimated cardinal Ts were 1 °C for the base T, 30 °C for the optimum T, and 40.8 °C for the ceiling T. The salt and drought tolerance threshold values were 257 mM NaCl and − 1.2 MPa for SG and 247 mM NaCl and − 1 MPa for the seedling growth, respectively, suggesting that the seedling growth was more sensitive than SG under both salt and drought stresses in Eruca. In addition, the maximum SG and seedling growth were observed at pH 7 and burial depth 1.9 cm. In general, the models used in this study could describe well the response of Eruca SG under different levels of environmental factors and also their parameters could easily be used in Eruca SG simulation models. This information also could help us to better manage the production of this plant under stressful conditions and/or to determine its geographic range expansion in the world.