Southern Limit Of Distribution Research Articles

Influence of climate change on flowering phenology of Yoshino cherry at its southern distribution limit.

To clarify the influence of climate change on the flowering phenology of Yoshino cherry at its southern distribution limit, we examined the relationship between cold exposure for endodormancy release (chilling requirement) and heat requirement for bud growth on Hachijojima Island, Japan, from 1948 to 2024. Cold exposure and heat requirement had a significant relationship approximated by linear or log-linear functions. In years with less cold exposure, the first flowering dates were much later than normal, in accordance with the higher heat requirement. Our results indicate that the variation in the balance between cold exposure and heat requirement depending on the pattern of annual air temperature change is likely to vary the first flowering date greatly at the distribution limit of Yoshino cherry.

A taxonomic revision of Tillandsia (Tillandsioideae, Bromeliaceae) from Uruguay

A taxonomic revision of the genus Tillandsia (Bromeliaceae) from Uruguay is here presented based on the examination of herbarium specimens, field surveys, and an extensive analysis of taxonomic literature. Nineteen taxa are recognized: 17 species, one subspecies, and one natural hybrid. A new identification key for all the species of Tillandsia occurring in the country is presented. Data on nomenclature (correct names, types, synonyms used in Uruguay) are provided for each taxon, along with detailed morphological descriptions, vernacular names, distribution and habitat information (global and local), phenology, conservation assessment, diagnostic characters, and nomenclatural and taxonomical notes. This information is complemented with images, illustrations of floral parts, maps showing species distribution in Uruguay, and a phenological diagram. Tillandsia arequitae and T. uruguayensis are endemic to Uruguay, and five species reach their southern limit of distribution in the country (T. bandensis, T. geminiflora, T. ixioides, T. loliacea, and T. stricta). Tillandsia capillaris and T. virescens are registered as exotic and subspontaneous. A new natural hybrid between T. aëranthos and T. recurvifolia is described. Three species previously cited for Uruguay (T. crocata, T. lorentziana, and T. streptocarpa) are excluded or cited as doubtful species for the country. Tillandsia bicolor is discarded as a synonym of T. aëranthos, and T. dianthoidea is considered for the first time as an illegitimate superfluous name. Lectotypes for Tillandsia arequitae (second-step), T. capillaris, T. circinalis (second-step), T. cordobensis (second-step), T. duratii, T. geminiflora (second-step), T. hieronymi (second-step), T. ixioides, T. microxiphion, T. recurvata, T. stricta (second-step), T. tricholepis (second-step) and T. virescens, as well as an epitype for T. aëranthos are designated.

Habitat Use of the Hen Harrier (Circus cyaneus) during the Breeding Season in Spain

Raptors usually show use for a particular habitat to settle during the breeding period. In this study, we aimed to study the habitat use of the breeding populations of Hen Harrier (Circus cyaneus) in Spain, which represents the southern distribution limit of the species. We used GPS/GSM data from 17 individuals during 21 breeding periods. We analysed space and habitat use using 95% kernel density estimators to obtain home ranges during the breeding period. Then, we performed a third-order habitat selection analysis, extracting the percentage of locations on each land use and comparing it with random points generated within each home range. Furthermore, an Agricultural Use Index was created to classify harriers with agricultural or forest use depending on habitat. The results showed that harriers had a general use for agricultural land and, to a lesser extent, habitats with natural vegetation scrubs. Nest location was key to habitat selection, and these uses varied with latitude, with natural vegetation being chosen as the nesting habitat in the north and agricultural fields in the south, probably due to the availability of habitats in different parts of Iberia.

Temperature dependence of feeding activity in the invasive freshwater snail Pomacea canaliculata: implications for its response to climate warming

AbstractPomacea canaliculata, a freshwater snail native to South America, has become widely naturalised in many countries, causing serious damage to agricultural and native ecosystems. Although climate warming is likely to expand the distribution of this species, limited information exists regarding its impact on feeding activities. In this study, we examined the effects of temperature on the feeding activity of this species and estimated the impact of climate warming on its feeding potential. The feeding activity was determined by measuring the amount of standard food (Japanese mustard spinach, komatsuna) ingested at different temperatures. It tended to increase from 15 to 25 °C but became almost stable from 25 to 35 °C. The respiration rate determined by the O2 consumption rate, showed a similar response to temperature. Based on these findings, we constructed a simple model to estimate the relative feeding activity using climatic data recorded at meteorological stations throughout the Japanese Archipelago. The model estimated that, with warming of + 2 °C, annual feeding potential (relative value) increased by 21.1% at the present northern distribution limit of this species. The effect of warming on percentage increase in feeding potential was estimated to be smaller at the southern distribution limit (9.9%), although the absolute feeding potential was larger than that at the northern sites. The model also suggested that if this species expanded its northern distribution range as a result of climate warming, it would have a high feeding potential comparable to that of the southern regions.

Diversity and distribution of the lichen genus Umbilicaria in the Argentine Islands–Kyiv Peninsula region, the maritime Antarctic

This paper documents the occurrence of the genus Umbilicaria in the Argentine Islands–Kyiv Peninsula region of the Graham Coast in the maritime Antarctic. The presence of seven Umbilicaria species (U. africana, U. antarctica, U. aprina, U. decussata, U. kappenii, U. nylanderiana and U. umbilicarioides) in the ice-free areas of the Argentine Islands–Kyiv Peninsula region were confirmed. The species of U. africana and U. aprina are documented from the studied region for the first time. This study moves the southern distribution limit of U. africana about 300 km to the south: to the Argentine Islands–Kyiv Peninsula region. The distribution maps of Umbilicaria species for the studied region and maritime Antarctica are prepared.

Seasonal Variability in Non-Structural Carbohydrate Content of Warm-Adapted Zostera noltei and Zostera marina Populations

Non-structural carbohydrates (NSCs) are energetic compounds that can be accumulated in tissues and mobilized during periods of unfavorable conditions to maintain the biological functions of plants. The balance of these biochemical compounds is controlled by environmental factors such as temperature and irradiance. Zostera noltei and Zostera marina find one of their southern distribution limits in southern Spain, where relatively high seawater temperatures are reached during summer (23–24 °C). To better understand the effects of elevated temperatures on the concentration of NSCs, we conducted a seasonal study at Cadiz Bay, representing warm-adapted populations of these species. Our results showed a bimodal pattern in both species, with the highest NSC content observed in December and June, followed by a depletion in March and August. In addition, the NSC content observed in the leaves of Z. noltei (71.26 ± 30.77 mg g−1 dry weight) was higher than in the rhizomes and roots (52.14 ± 38.86 mg g−1 DW). The observed patterns suggest that these species accumulated NSCs to cope with periods of unfavorable environmental conditions. We also suggest that the limited concentration of NSCs in Z. noltei rhizomes and roots indicates that this population may be suffering physiological stress.

Caught in transition: changes in brachyuran diversity following mangrove encroachment into saltmarshes at a southern distribution limit

Mangroves are expanding polewards due to global change, often encroaching into adjacent temperate saltmarshes. In both vegetated ecosystems, brachyurans are responsible for ecological processes and functions such as nutrient cycling and sediment bioturbation. South African mangroves occur at a latitudinal limit and are establishing further south due to past planting events and global change, making these ideal study systems for the effects of mangrove expansion and encroachment. Here, we investigated the effect of mangrove encroachment on brachyuran community composition at two saltmarsh sites with planted mangrove stands of different ages. Transects were laid perpendicular to each estuary where three habitat types were demarcated (mangrove, ecotone, saltmarsh). Sediment samples were collected for analyses and quadrats were used to measure pneumatophore density, saltmarsh cover, and brachyuran abundance and diversity. We found that brachyuran community structure at each site has significantly changed over seven years, with two mangrove-associated fiddler crab species, Tubuca urvillei and Paraleptuca chlorophthalmus, now recorded at the younger planted site, indicating a new southern distributional limit. Community structure was also significantly different amongst habitat types (p < 0.05) with Parasesarma catenatum dominating saltmarshes while Danielella edwardsii was more prominent in mangroves. However, community composition did not differ significantly between the two (differently aged) sites (p > 0.05). Pneumatophore density had a proportional relationship with crab abundance, diversity and richness, while saltmarsh cover had an inversely proportional relationship with crab abundance, diversity and richness. It is likely that as mangroves continue to expand into saltmarshes, more mangrove-associated species will move into saltmarshes, potentially altering ecosystem processes in this unique habitat.

Twenty years of population dynamics in European beech-oak forest at their rear range margin anticipate changes in its structure and composition

There is an increasing interest in restoring degraded forests, which occupy half of the forest areas. Among the forms of restoration, passive restoration, which involves the elimination of degrading factors and the free evolution of natural dynamics by applying minimal or no management, is gaining attention. Natural dynamics is difficult to predict due to the influence of multiple interacting factors such as climatic and edaphic conditions, composition and abundance of species, and the successional character of these species. Here, we study the natural dynamics of a mixed forest located in central Spain, which maintained an open forest structure, due to intensive use, until grazing and cutting were banned in the 1960s. The most frequent woody species in this forest are Fagus sylvatica, Quercus petraea, Quercus pyrenaica, Ilex aquifolium, Sorbus aucuparia, Sorbus aria and Prunus avium, with contrasting shade and drought tolerance. These species are common in temperate European deciduous forest and are found here near their southern distribution limit, except for Q. pyrenaica. In order to analyze forest dynamics and composition, three inventories were carried out in 1994, 2005 and 2015. Our results show that, despite the Mediterranean influence, the natural dynamics of this forest has been mainly determined by different levels of shade tolerance. After the abandonment of grazing and cutting, Q. pyrenaica expanded rapidly due to its lower shade tolerance, whereas after canopy closure and forest densification, shade-tolerant species gained ground, particularly F. sylvatica, despite its lower drought and late-frost tolerance. If the current dynamics continue, F. sylvatica will overtake the rest of the species, which will be relegated to sites with shallow soils and steep slopes. Simultaneously, all the multi-centennial beech trees, which are undergoing a rapid mortality and decline process, will disappear.

Massive barnacle recruitment on the Gulf of St. Lawrence coast of Nova Scotia (Canada) in 2024 linked to increased sea surface temperature.

With the ongoing climate and oceanographic change, an increasing number of studies are reporting dramatic population losses caused by thermal extremes in intertidal habitats. Under moderate warming, however, populations can fare better in places where species normally experienced suboptimal temperatures. This article reports the massive recruitment of the barnacle Semibalanus balanoides on the Gulf of St. Lawrence coast of Nova Scotia (Canada) in 2024. As recruits appear mostly during May in this region, coastal sea surface temperature (SST) in April is critical for the ecological performance of larvae, as they are pelagic and live in the water column for weeks before intertidal settlement. Thus, a study that spanned 12 years (2005 to 2016) on this coast found that annual barnacle recruitment was positively correlated to April SST. In April 2024, coastal SST was 116% higher than for the same month averaged over those 12 years (4.1 vs. 1.9 °C). This SST spike was followed by an elevated recruitment that was 111% higher than the average for those 12 years (1,278 vs. 607 recruits dm-2). Overall for the studied years, the amount of variation in annual barnacle recruitment statistically explained by April SST was 51%. While the southern distribution limit of S. balanoides has moved northwards in recent decades due to lethal warming, our results support the notion of improving reproductive success with seawater warming on colder northern shores.

The intra-annual rhythm of Pinus sylvestris growth-climate responses under a warming climate at its southern distribution limits

Warming climate has posed a pressing threat on boreal forests and an improved understanding of the intra-annual pattern of climatic influences on the tree growth can help interpret the response of boreal forest to climate change. Here, we systematically examined the growth-climate relationship, cambium phenology, and xylem cell dynamics of Pinus sylvestris (PS) to disentangle its intra-annual growth rhythm to external drivers and internal physiological process at its southern distribution limits across Eurasia. We showed the intra-annual rhythm of PS to temperature and precipitation is anti-phase and synchronous at its southern limits. Temperature had both promoting and inhibiting effects on PS growth over growing season where the growth-temperature response gradually changed from positive in winter-early spring to negative in late spring-summer, and then returned to positive in autumn. Precipitation enhanced PS growth in late spring-summer. Temperature dominated the intra-annual growth-climate response rhythm, and the response shifted to negative when temperature reached 13.72 °C in spring-summer and shifted to positive when temperature reached 17.41 °C in summer-autumn in Shenyang, respectively. A high water demand for the rapid earlywood cell formation in spring-summer and heat requirement for cambial cell division in summer-autumn caused the shifts in growth-temperature response, respectively. Climatic warming advances and prolongs the time of summer water availability that limits PS growth at its southern distribution limits and thus warming climate may pose a greater threat to the southern population of PS.

Modeling and updating the occurrence of Aedes aegypti in its southern limit of distribution in South America

Aedes aegypti is the main vector of arboviruses in the world. This mosquito species is distributed from tropical to temperate regions. In Argentina, it has been reported in 20 out of 23 provinces and reaches its southernmost distribution in the world. Its distribution and persistence are affected by meteorological, demographic and environmental factors, such as temperature, precipitation, and population. The aim of this study was to update and model the occurrence of Aedes aegypti in its southern limit of distribution in Argentina. To this end, a total of 37 sites were inspected in La Pampa and Río Negro provinces. Generalized Linear Models were used to explain the occurrence of Aedes aegypti based on meteorological, environmental and demographic variables. Aedes aegypti was found in 11 cities of La Pampa province where it had not been previously reported, but was not found in any of the cities evaluated in Río Negro province. The averaged model explaining the occurrence of Aedes aegypti included the minimum temperature, precipitation and interactions between maximum temperature and precipitation as explanatory variables. Although precipitation was statistically significant, other factors such as minimum temperature are also important in modeling the occurrence of Aedes aegypti in its southernmost distribution limit.

Ecophysiological Behavior of Fagus sylvatica L. Growing at Its Southern Distribution Limit: Insights for Understanding the Fate of the European Beech under Warmer and Dryer Growth Conditions

In the last 20 years, a significant mortality of Fagus sylvatica L. (European beech) has been documented in central and northern European forests. Surprisingly, no beech die off occurred at the southern limit. This fact leads us to hypothesize that European beech populations growing at the southern limit of the distribution might have a significant phenotypic plasticity to better cope with low water availability and rising temperatures. To check this hypothesis, we evaluated the ecophysiological behavior of F. sylvatica growing along an altitudinal transect in Calabria (Italy). We selected three study sites (750 m a.s.l., 976 m a.sl. m a.s.l., 1450 m a.s.l.) showing narrow ranges of temperature, rainfall and air humidity. Trees growing at 976 m a.s.l. showed the highest stomatal conductance values during the entire experimental period. The lowest gas exchange and highest leaf mass area were recorded in plants growing at 750 m a.s.l. In the European beech growing at 1450 m a.s.l., higher vessel density, lower mean vessel diameter and higher vessel grouping index values were recorded. Overall, our results highlighted that the measured populations show a considerable phenotypic plasticity leading them to adjust anatomical and physiological traits in response to narrow ranges of environmental parameters. Despite that, the distribution of F. sylvatica seems to be limited to areas with a growing season rainfall of at least 400 mm and vapor pressure deficit (VPD) values < 3 kPa, which may represent the main environmental thresholds which strongly limit the beech growth and, therefore, influence the ability of this species to cope with future environmental conditions.

Drivers and dynamics of foliar senescence in temperate deciduous forest trees at their southern limit of distribution in Europe

Research on autumn phenology is very important for understanding and simulating the future growth of temperate deciduous forests. This is especially needed at the southern edge of the temperate zone, where climate change impacts are particularly intense. We studied foliar senescence timing for mature stands of Fagus sylvatica L., Populus tremula L., Betula pendula Roth, Quercus petraea (Matt.) Liebl. and Quercus robur L. at the southern edge of European temperate forests. First, we analysed long-term series (1997–2019) of senescence timing in southern France. Then, we compared a more detailed four-year dataset (2017–2020) of senescence dynamics and its correlations to meteorological conditions for stands in northern Spain and Belgium, with the latter area representing the core distribution of the species. In contrast to other temperate regions, no significant delaying trends in the timing of autumn phenology were detected in the long-term in southern temperate forests, which were characterized by high inter-site variability. In the short-term (4 years), species-specific correlations with meteorological conditions were found, with, for example, the senescence of F. sylvatica being affected by temperature while the senescence of Q. petraea was affected by both temperature and precipitation. Autumn dynamics differed between the core and southern areas of distribution of the species in the study period. In particular, while late season conditions affected senescence timing in the southern populations, the senescence dynamics of the core populations was mostly influenced by the legacy of spring – early summer conditions and, overall, more affected by precipitation than southern populations. Our data fill important knowledge gaps on the functioning of temperate deciduous forests at the southern limit of distribution in Europe.

Development of nuclear microsatellite markers in Yerba mate (Ilex paraguariensis A. St. Hil.) from whole-genome sequence data

AbstractIlex paraguariensis A. St.-Hil. (yerba mate) (Aquifoliaceae Bercht. & J. Presl) is a plant species with great economic and cultural importance because its leaves are processed and ground to make infusions like mate or tereré. The species is distributed in a continuous area that includes Southern Brazil and part of Paraguay and Argentina. Uruguay represents the Southern distribution limit of the species, where small populations can be found as part of ravine forests. Although there are previous reports of molecular markers for this and other species in the genus, the available markers were not informative enough to represent the intra- and interpopulation genetic diversity in marginal Uruguayan populations. In this study, we developed highly informative polymorphic microsatellite markers to be used in genetic studies in I. paraguariensis. Markers were identified in contigs from the genome sequence of two individuals and then tested for amplification and polymorphism content in a diverse panel. Markers which passed these filters were tested on populations from Uruguay. They detected higher diversity within populations (in terms of number of alleles and heterozygosity) than previously reported, and levels of heterozygosity similar to those reported for two Brazilian populations. This subset of seven markers were successfully multiplexed, substantially reducing the costs of the analysis. Combined with previously reported nuclear and plastid markers, they can be used to evaluate the genetic diversity of rear-edge populations, identify genotypes for paternity studies and provide relevant information for the conservation and management of germplasm.

Sensitivity of Eurasian Rear-Edge Birch Populations to Regional Climate and Local Hydrological Conditions

South rear-edge populations of widely distributed temperate and boreal tree species such as birches (Betula pubescens and Betula pendula) are considered particularly vulnerable to climate warming, and at the same time, they constitute genetic reservoirs of drought-adapted ecotypes. Here, we compared radial growth patterns and responses to climate, river, or reservoir flows and a drought index of rear-edge (southernmost) populations (Toledo Mountains, central-southern Spain) with populations located in northern Spain of B. pubescens and B. pendula. Then, we performed a comparative analysis across Europe of B. pendula populations. The main climatic constraint of birch growth was a high summer water deficit, although the effect of local hydrological conditions was particularly important in rear-edge populations. We found declining growth trends in rear-edge stands dating from the early 21st century, related to decreasing water availability and increasing aridity. Our results also suggested distinct growth patterns and climate-growth associations of B. pendula across Europe that show how populations further south and in warmer locations were more sensitive to drought stress. Drought-induced growth decline can be exacerbated by local human land uses, leading to reduced river inflow, thus endangering birch populations at their southern distribution limit. Protection of threatened rear-edge birch populations requires adequate management of local water resources.

Whiting (Merlangius merlangus) Grows Slower and Smaller in the Adriatic Sea: New Insights from a Comparison of Two Populations with a Time Interval of 30 Years

Nowadays, overexploitation and climate change are among the major threats to fish production all over the world. In this study, we focused our attention on the Adriatic Sea (AS), a shallow semi-enclosed sub-basin showing the highest exploitation level and warming trend over the last decades within the Mediterranean Sea. We investigated the life history traits and population dynamics of the cold-water species whiting (Merlangius merlangus, Gadidae) 30 years apart, which is one of the main commercial species in the Northern AS. The AS represents its southern limit of distribution, in accordance with the thermal preference of this cold-water species. Fish samples were collected monthly using a commercial bottom trawl within the periods 1990–1991 and 2020–2021. The historical comparison highlighted a recent reduction in large specimens (>25 cm total length, TL), which was not associated with trunked age structures, therefore indicating a decrease in growth performance over a period of 30 years (L∞90–91 = 29.5 cm TL; L∞20–21 = 22.8 cm TL). The current size at first sexual maturity was achieved within the first year of life, at around 16 cm TL for males and 17 cm TL for females. In the AS, whiting spawns in batches from December to March, showing a reproductive investment (gonadosomatic index) one order of magnitude higher in females than in males. Potential fecundity (F) ranged from 46,144 to 424,298, with it being heavily dependent on fish size. We hypothesize that the decreased growth performance might be related to a metabolic constraint, possibly related to the increased temperature and its consequences. Moreover, considering the detrimental effects of size reduction on reproductive potential, these findings suggest a potential endangerment situation for the long-term maintenance of whiting and cold-related species in the AS, which should be accounted for in setting management strategies.

Sea Cucumbers (Echinodermata: Holothuroidea) from Santa Catarina Coast, Southern Brazil, with Notes on Their Abundance and Spatial Distribution

Sea cucumbers are important components of marine ecosystems and also highly valued commodities in Asian markets because of their use in oriental cuisine and traditional medicine. The increasing demand for sea cucumbers has elicited intense worldwide capture and many species are currently considered vulnerable or threatened due to overexploitation. The coast of Santa Catarina State (South Brazil) is the distribution limit of several tropical species in the Western South Atlantic, but little is known about the sea cucumber species occurring in this region. This study aimed to identify the holothuroids occurring in the littoral of Santa Catarina State and to survey their abundance and distribution along the coast. Visual and tactile inspections were carried out following transects in the intertidal zone of 11 locations and subtidal surveys were carried out by diving in two sites. The surveys indicated the occurrence of three species: Parathyone braziliensis in the intertidal zone, Holothuria (Halodeima) grisea in the intertidal as well as in the sublittoral, and Isostichopus badionotus in the sublittoral. The most abundant species was H. (H.) grisea, which occurred in several sites along the coast, up to Lat. 28° 01′ 07.67″ S, its likely southern limit of distribution. The species I. badionotus was for the first time recorded to occur on the continental coast of the State, expanding its area of distribution. This study registers for the first time the occurrence of P. braziliensis in southern Brazil, expanding its limit of geographic distribution.

Citizen science aids the quantification of the distribution and prediction of present and future temporal variation in habitat suitability at species’ range edges

Citizen science programs are effective methods to collect large volumes of data to assist researchers in monitoring ecological environments. As species shift their distributions globally due to climate change, the use of citizen science data to detect these shifts is increasing. Using targeted citizen science programs to collect data on these species could provide information on range edges to inform species distribution modelling. Currently, species distribution models (SDMs) often rely on large data repositories that may lack observations, and hence ability, to detect changes at the range edge. Here, we developed a SDM to compare traditional data repository observations with targeted citizen science data at the southern distribution limit of two recreationally important marine fish in Tasmania, Australia to investigate the potential change in spatial predictions at their range edge. The SDM using the targeted citizen science data in addition to traditional observation data improved the representation of species by 2.3 and 52.7% and increased the southern distribution of the species by 277 and 438 km, for snapper and King George whiting, respectively. Future (centred around 2050 under IPCC RCP 8.5) habitat suitability was predicted to increase more over the winter season, with implications for species overwintering and persistence of populations. The use of citizen science data allowed for the modelling of historical and future change for two range-extending species, an outcome possible due to the collaboration of two citizen science programs that collected observational data on the target species. Species range shifts will require ongoing monitoring and we have demonstrated that complimentary citizen science initiatives are effective in capturing occurrences of species at their range edge. Increasing collaboration between programs may further increase data collection efforts and provide the knowledge to create a hub for these data to be used more efficiently in the future.

Dieback of the cushion plant Silene acaulis at its southern limit of distribution in the Apennines

Dieback of the cushion plant Silene acaulis at its southern limit of distribution in the Apennines

Phenotypic specialization of the pea aphid in its southern limit of distribution

The success of biological invasions ultimately relies on phenotypic traits of the invasive species. Aphids, which include many important pests worldwide, may have been successful invading new environments partly because they can maximize reproductive output by becoming parthenogenetic and losing the sexual phase of their reproductive cycle. However, invasive populations of aphids invading wide ranges can face contrasting environmental conditions and requiring different phenotypic strategies. Besides transitions in their reproductive cycle, it is only partially known which phenotypic traits might be associated to the invasion success of aphid populations in extended novel ranges. Here, we used four genotypes of the pea aphid Acyrthosiphon pisum from two localities in Chile to test for phenotypic specialization that might explain their establishment and spread in habitats exhibiting contrasting environmental conditions. We show that lineages living at a higher latitude with low temperatures show, in addition to facultative sexual reproduction, smaller body sizes, lower metabolic rates and a higher tolerance to the cold than the obligate asexual lineages living in a mild weather, at the expense of fecundity. Conversely, at higher temperatures only asexual lineages were found, which exhibit larger body sizes, higher reproductive outputs and consequently enhanced demographic ability. As a result, in conjunction with the reproductive mode, lineage specialization in physiological and life-history traits could be taken into account as an important strategy for populations of pea aphid to effectively invade extended novel ranges comprising different climatic conditions.