Novel spatial analysis methods reveal scale‐dependent spread and infer limiting factors of invasion by Sahara mustard

Multiple scale‐dependent ecological processes influence species distributions. Uncovering these drivers of dynamic range boundaries can provide fundamental ecological insights and vital knowledge for species management. We develop a transferable methodology that uses widely available data and tools to determine critical scales in range expansion and to infer dominating scale‐dependent forces that influence spread. We divide a focal geographic region into different sized square cells, representing different spatial scales. We then used herbarium records to determine the species' occupancy of cells at each spatial scale. We calculated the growth in cell occupancy across scales to infer the scale dependent expansion rate. This is the first time such a ‘box‐counting’ method is used to study range expansion. We coupled this multi‐scale analysis with species distribution models to determine the range and spatial scales where suitable climate allows the species to spread, and where other factors may be influencing the expansion. We demonstrate our methodology by assessing the spread of invasive Sahara mustard in North America. We detect critical scales where its spread is limited (100–500 km) or unconstrained (5–50 km) by climatic variables. Using climate‐based models to assess the similarity of climate envelopes in its native and invaded range, we find that the climate in the invaded range generally predicts the native distribution, suggesting that either there has been little local adaptation to climate occurring since introduction or the biological interaction experienced in the invaded range has not driven the species to occupy climatic conditions much different from its native range. Our novel method can be broadly utilized in other studies to generate critical insights into the scale dependency of different ecological drivers that influence the spread and distribution limits, as well as to help parameterizing predictions of future spread, and thus inform management decisions.     

The taxonomy and phytogeography of bracken—a review

Some of the biological problems presented by bracken, Pteridium aquilinum (L.) Kuhn, are posed. Its taxonomic position within the Pteridophyta and the delimitation of entities within the genus are discussed on the basis of morphological and cytological evidence. The geographical ranges of the various brackens world-wide are described and mapped in outline, and emphasis placed on reviewing the natural ecological role of bracken in plant communities throughout the world. Further geographic areas where taxonomic investigation of Pteridium is most needed are indicated, and evidence of the reproductive, dispersal, establishment, colonizing ability and vegetative persistence of bracken is reviewed. Its palaeobiological spread, with associated vegetational history, and the effects on this of anthropogenic influences—better known than are comparable details for any other pteridophyte—are detailed, and the present magnitude of the resulting bracken problem in Britain (and especially in upland Britain) indicated.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

Bracken invasion in Scotland

Summary

For the first time, estimates of the amount of sparse bracken on open land, the amounts of bracken under tree cover and the amounts in linear features are made. These data were used to predict the areas of Scotland most at risk from invasion from patch edges, from sparse patches becoming dense and from invasion along linear features. Computer models show that if the climate warms over the next 50 yr as predicted, then bracken growth will increase and the amount of hill land suitable for invasion by bracken will increase. On some sites where bracken is being controlled, it is reduced in cover and other ground vegetation is becoming established. However, we cannot ensure that when bracken is controlled that the control is always effective and that appropriate vegetation will become established. Some generalized rules have been derived to help in the selection of sites likely to show the most benefit from control treatment.     

The heat is on: frequent high intensity fire in bracken (<Emphasis Type="Italic">Pteridium aquilinum</Emphasis>) drives mortality of the sprouting tree <Emphasis Type="Italic">Protea</Emphasis><Emphasis Type="Italic">caffra</Emphasis> in temperate grasslands

We examined the effect of fire frequency and intensity on a Protea caffra tree population in the temperate montane grasslands of north-western KwaZulu-Natal, South Africa. We assessed the effect of fire by comparing the population structure of the resprouter P. caffra in discrete bracken (Pteridium aquilinum) patches with that in the surrounding grassland matrix. Fuel biomass did not differ between grassland and bracken, but bracken fuel was significantly drier than grass. Above-ground fire temperatures and fireline intensity, measured by P. caffra char height, were significantly higher in the bracken habitat. Forty-two percent of the P. caffra population in grassland and in bracken persisted by coppice resprouts, having lost their original stem to fire damage. Exposure to higher intensity bracken fire suppressed P. caffra regeneration and caused greater adult mortality compared with trees in grassland. Consequently, the P. caffra population in bracken was skewed towards old age with most trees severely fire damaged. The high incidence of small trees in grassland indicates that a regular fire interval of 2–3 years does not negatively affect regeneration of P. caffra. However, in bracken patches regular high intensity fires cause high mortality among all P. caffra size classes and will ultimately result in local extinction. Bracken thus has the potential to significantly alter tree–grass interactions in these montane grasslands.     

Bracken-induced increase in soil P availability,along with its high P acquisition efficiency,enables it to invade P-deficient meadows

欧洲蕨入侵提高了土壤磷有效性和高磷获取效率 对于欧洲蕨(Pteridium aquilinum)入侵荒地后土壤化学的变化,已经有了相关的研究,但是对于欧洲蕨入侵草地的研究却比较缺乏。本研究探讨了欧洲蕨入侵缺磷草地是否会改变土壤养分资源库,以及与土壤过程和欧洲蕨营养相关的机制。此外,还研究了欧洲蕨入侵前后群落组成对土壤化学差异的反应。在蕨菜生物量高峰期间,我们进行了土壤和植物取样以及植被调查。数据分析包括方差分析(ANOVA)和典型对应分析(CCA)。研究结果表明,欧洲蕨入侵提高了土壤磷有效性、土壤有机碳浓度以及碳氮比、碳硫比和氮硫比,同时降低了铁和钴的浓度。欧洲蕨羽片富含磷,根茎富含钾,而羽片 和根茎的氮磷比很低。典型对应分析(CCA)显示了与磷和钾有效性相关的常见草地植物物种的不同丰度模式。在磷有效性极低的条件下,绒毛草(Holcus lanatus)表现出竞争优势。欧洲蕨通过促进铁和铝的浸出提高了磷有效性。从土壤资源生态位的角度来看,欧洲蕨通过提高自身生长所需的磷有效性并且增加对其他物种的氮限制,从而获得了竞争优势。欧洲蕨提高土壤磷有效性的能力,以及其高磷获取效率背后的生理机制,似乎将欧洲蕨与其他竞争生态策略的物种区分开来,因为其他物种的生长主要局限于营养丰富的环境,因此欧洲蕨更容易入侵缺磷的草地。     

Control of bracken

Mechanical methods of bracken control are now being replaced by the use of chemicals. Trials indicate the value of asulam and glyphosate in reducing frond numbers of bracken when sprayed in late July or early August in the west of Scotland. There is still a need to find a bracken eradicant chemical rather than a control chemical. Research is also required to investigate the results of removing the bracken cover on the ecology of the treated areas. The effects of sudden exposure of the hitherto protected grass to extremes of climate are not known.     

Climatic niche breadth determines the response of bumblebees (<Emphasis Type="Italic">Bombus</Emphasis> spp.) to climate warming in mountain areas of the Northern Iberian Peninsula

Studies examining species range shifts in the face of climate change have consistently found that response patterns are complex and varied, suggesting that ecological traits might be affecting species response. However, knowledge of how the traits of a species determine its response to climate change is still poorly understood. Here we investigate the role of species-specific climate niche breadth in forecasting bumblebee (Bombus spp.) responses to regional climate warming in the Cantabrian Range (north-western Iberian Peninsula). Climate niche breadth was defined using known data for occurrences of specific species at their continental (i.e., European) scale of distribution. For each bumblebee species, climate niche breadth was found to be related to (1) the elevational range shifts of species between their historical (1988–1989) and recent (2007–2009) distribution and (2) the variation in the climatic conditions of the localities they inhabited (i.e., the local climate space) between both study periods. Our results show a strong relationship between climate niche breadth, particularly thermal niche breadth, and the response of bumblebee species to climate warming, but only when this response was determined as variations in local climate space. The main conclusions of our work are thus twofold. First, variations in the climatic conditions underlying range shifts are useful in making accurate assessments of the impact of climate change on species distributions. Second, climate niche breadth is a particularly informative ecological trait for forecasting variations in species responses to climate change.     

Dispersal and species’ responses to climate change

Dispersal is fundamental in determining biodiversity responses to rapid climate change, but recently acquired ecological and evolutionary knowledge is seldom accounted for in either predictive methods or conservation planning. We emphasise the accumulating evidence for direct and indirect impacts of climate change on dispersal. Additionally, evolutionary theory predicts increases in dispersal at expanding range margins, and this has been observed in a number of species. This multitude of ecological and evolutionary processes is likely to lead to complex responses of dispersal to climate change. As a result, improvement of models of species’ range changes will require greater realism in the representation of dispersal. Placing dispersal at the heart of our thinking will facilitate development of conservation strategies that are resilient to climate change, including landscape management and assisted colonisation. Synthesis This article seeks synthesis across the fields of dispersal ecology and evolution, species distribution modelling and conservation biology. Increasing effort focuses on understanding how dispersal influences species' responses to climate change. Importantly, though perhaps not broadly widely‐recognised, species' dispersal characteristics are themselves likely to alter during rapid climate change. We compile evidence for direct and indirect influences that climate change may have on dispersal, some ecological and others evolutionary. We emphasise the need for predictive modelling to account for this dispersal realism and highlight the need for conservation to make better use of our existing knowledge related to dispersal.     

The history and ethnobotany of bracken

The fossil record shows that Pteridium has been present in all the temperate stages of the Quaternary, but that for most of this period it formed a component of the herb layer of deciduous woodlands. Bracken did not begin to reach its present abundance and importance until widespread forest clearance began with the arrival of Neolithic man about 5000 years ago. Some of the factors that have played a role in the history of bracken are illustrated by a discussion of the spread of bracken in Scotland that occurred contemporaneously with the change from cattle- to sheep-farming in the 18th century. It is shown that man has played a dominant role in influencing the spread of bracken in Scotland, but that bracken was considered of great positive economic value. The ethnobotany of bracken is discussed, and it is suggested that it was once an important source of potash for the glass, soap and bleaching industries. The use of the plant as fuel, thatch, litter, compost and food, and for medicinal purposes is considered, and mention is made of various minor uses of the plant. Unfortunately it is difficult to estimate the amount of bracken consumed by these various employments, but it is suggested that the increased abundance of the plant may have been exaggerated because what was once a useful resource has now become a pest.     

Species’ traits as predictors of range shifts under contemporary climate change: A review and meta‐analysis

A growing body of literature seeks to explain variation in range shifts using species’ ecological and life‐history traits, with expectations that shifts should be greater in species with greater dispersal ability, reproductive potential, and ecological generalization. Despite strong theoretical support for species’ traits as predictors of range shifts, empirical evidence from contemporary range shift studies remains limited in extent and consensus. We conducted the first comprehensive review of species’ traits as predictors of range shifts, collecting results from 51 studies across multiple taxa encompassing over 11,000 species’ responses for 54 assemblages of taxonomically related species occurring together in space. We used studies of assemblages that directly compared geographic distributions sampled in the 20th century prior to climate change with resurveys of distributions after contemporary climate change and then tested whether species traits accounted for heterogeneity in range shifts. We performed a formal meta‐analysis on study‐level effects of body size, fecundity, diet breadth, habitat breadth, and historic range limit as predictors of range shifts for a subset of 21 studies of 26 assemblages with sufficient data. Range shifts were consistent with predictions based on habitat breadth and historic range limit. However, body size, fecundity, and diet breadth showed no significant effect on range shifts across studies, and multiple studies reported significant relationships that contradicted predictions. Current understanding of species’ traits as predictors of range shifts is limited, and standardized study is needed for traits to be valid indicators of vulnerability in assessments of climate change impacts.     

The ecological status of bracken

Information about the morphology of bracken ( Pteridium aquilinum (L.) Kuhn var. aquilinum ) and the chief nutrients in the frond at different times of the year introduce an account of litter production and its accumulation in relation to the behaviour of frond, root and rhizome systems.
Where litter gain exceeds loss there is a correlation between the thickness and/or kind of litter and the level of the root and rhizome systems in relation to the mineral soil surface: with increase of litter the bracken becomes progressively more dependent for its physical and chemical soil environment on its own débris and less on the underlying mineral soil. An example of the limit of complete dependence has not been examined, but degeneration of the community can take place before that stage is reached.
From a review of the chief factors affecting bracken the conclusion is reached that the woodland habitat is both favourable and restrictive: in it bracken is in equilibrium with its environment, at a high social status. The relationship with other plant communities depends largely on the degree of human interference to which each is subject. Dominant bracken when left alone, and where gain of litter exceeds loss, becomes the victim of its own success; local degeneration opens the way for entry by other species.     

Phenotypic traits of the Mediterranean <Emphasis Type="Italic">Phragmites australis</Emphasis> M1 lineage: differences between the native and introduced ranges

The environmental conditions in the new ranges of introduced plant species are often different from the conditions in their native ranges, and invasive plant species have been assumed to adapt to different environmental conditions by rapid ecological evolution in the invasive range after the introduction. Another interpretation of the change in plant traits after their introduction, however, is ecological fitting, which is based on the inherently high phenotypic plasticity of the species rather than on evolution. The Mediterranean haplotype M1 lineage of the wetland grass Phragmites australis was introduced to the coastal wetlands along the Gulf Coast of North America, where it is exposed to a different climate compared to its original range. The climate in the native range is arid or temperate with dry and hot summers, whereas the climate in the introduced range is warmer and has a higher and more uniform precipitation than that in the native range. This warmer and more humid environment is likely to pose different selection pressures to the plants in the introduced range and thus cause rapid evolutionary change and phenotypic differentiation in the introduced range. Here, we compared phenotypic traits of the M1 lineage from the native and introduced ranges in a common garden experiment to study the processes assisting the successful spread in the introduced range. Overall, the native and introduced groups were similar, but we detected a few phenotypic traits that diverged. Ecological fitting could be the fundamental mechanism by which the P. australis M1 lineage survives and spreads in the introduced Gulf Coast region. However, further research is needed to assess how the diverging traits observed in our study in Denmark (lower photosynthetic rates, lower chlorophylls concentration and higher leaf K concentration for the introduced than for the native genotypes) are expressed in the two ranges.     

Haemolymph Defense Capacity of the Neotropical sawfly <Emphasis Type="Italic">Aneugmenus merida</Emphasis> against Ant Predation

The defensive characteristics of the sawflies have received special attention due to the involvement of toxic compounds obtained from host plants. In this context, the haemolymph-based defense is one of the strategies known in sawflies to dissuade the attack of predators. Aneugmenus merida is a neotropical sawfly whose larvae are herbivorous on the toxic bracken fern Pteridium spp. The present study examines the defensive properties of the A. merida larval haemolymph and its possible link with the chemistry of its host plant. We report the behavior of the solitary hunter ant Odontomachus chelifer towards A. merida larvae under laboratory conditions. In addition, we studied the liquid intake behavior of the ants provided with solutions of crude haemolymph, bracken extracts, and its fractions. A. merida larvae showed a marked defensive capacity against the ants. The inhibition of the attack was observed during the stages of antennal contact and mandibular blow, suggesting that larval defensive capacity is due to factors present in the integument and haemolymph. Aqueous and methanolic fractions of haemolymph and bracken also deterred feeding. Although some common compounds were detected in the haemolymph and bracken fractions, they were in very small quantities, suggesting that they are not responsible for the bioactivity. Therefore, the hypothetical connection between the host plant chemistry and larval defensive capacity could not be evidenced. We suggest that the deterrent compounds present in the haemolymph and integument could be jointly acting in the sawfly’s defensive strategy.     

Are large‐scale distributional shifts of the blue‐winged macaw (Primolius maracana) related to climate change?

Aim To evaluate whether observed geographical shifts in the distribution of the blue‐winged macaw (Primolius maracana) are related to ongoing processes of global climate change. This species is vulnerable to extinction and has shown striking range retractions in recent decades, withdrawing broadly from southern portions of its historical distribution. Its range reduction has generally been attributed to the effects of habitat loss; however, as this species has also disappeared from large forested areas, consideration of other factors that may act in concert is merited. Location Historical distribution of the blue‐winged macaw in Brazil, eastern Paraguay and northern Argentina. Methods We used a correlative approach to test a hypothesis of causation of observed shifts by reduction of habitable areas mediated by climate change. We developed models of the ecological niche requirements of the blue‐winged macaw, based on point‐occurrence data and climate scenarios for pre‐1950 and post‐1950 periods, and tested model predictivity for anticipating geographical distributions within time periods. Then we projected each model to the other time period and compared distributions predicted under both climate scenarios to assess shifts of habitable areas across decades and to evaluate an explanation for observed range retractions. Results Differences between predicted distributions of the blue‐winged macaw over the twentieth century were, in general, minor and no change in suitability of landscapes was predicted across large areas of the species’ original range in different time periods. No tendency towards range retraction in the south was predicted, rather conditions in the southern part of the species’ range tended to show improvement for the species. Main conclusions Our test permitted elimination of climate change as a likely explanation for the observed shifts in the distribution of the blue‐winged macaw, and points rather to other causal explanations (e.g. changing regional land use, emerging diseases).     

Munroa argentina,a Grass of the South American Transition Zone,Survived the Andean Uplift,Aridification and Glaciations of the Quaternary

The South American Transition Zone (SATZ) is a biogeographic area in which not only orogeny (Andes uplift) and climate events (aridification) since the mid-Miocene, but also Quaternary glaciation cycles had an important impact on the evolutionary history of the local flora. To study this effect, we selected Munroa argentina, an annual grass distributed in the biogeographic provinces of Puna, Prepuna and Monte. We collected 152 individuals from 20 localities throughout the species’ range, ran genetic and demographic analyses, and applied ecological niche modeling. Phylogenetic and population genetic analyses based on cpDNA and AFLP data identified three phylogroups that correspond to the previously identified subregions within the SATZ. Molecular dating suggests that M. argentina has inhabited the SATZ since approximately 3.4 (4.2–1.2) Ma and paleomodels predict suitable climate in these areas during the Interglacial period and the Last Glacial Maximum. We conclude that the current distribution of M. argentina resulted from the fragmentation of its once continuous range and that climate oscillations promoted ecological differences that favored isolation by creating habitat discontinuity.     

Local adaptation and the evolution of species’ ranges under climate change

The potential impact of climate change on biodiversity is well documented. A well developed range of statistical methods currently exists that projects the possible future habitat of a species directly from the current climate and a species distribution. However, studies incorporating ecological and evolutionary processes remain limited. Here, we focus on the potential role that local adaptation to climate may play in driving the range dynamics of sessile organisms. Incorporating environmental adaptation into a stochastic simulation yields several new insights. Counter-intuitively, our simulation results suggest that species with broader ranges are not necessarily more robust to climate change. Instead, species with broader ranges can be more susceptible to extinction as locally adapted genotypes are often blocked from range shifting by the presence of cooler adapted genotypes that persist even when their optimum climate has left them behind. Interestingly, our results also suggest that it will not always be the cold-adapted phenotypes that drive polewards range expansion. Instead, range shifts may be driven by phenotypes conferring adaptation to conditions prevalent towards the centre of a species’ equilibrium distribution. This may have important consequences for the conservation method termed predictive provenancing. These initial results highlight the potential importance of local adaptation in determining how species will respond to climate change and we argue that this is an area requiring urgent theoretical and empirical attention.     

Eco‐evolution on the edge during climate change

We urgently need to predict species responses to climate change to minimize future biodiversity loss and ensure we do not waste limited resources on ineffective conservation strategies. Currently, most predictions of species responses to climate change ignore the potential for evolution. However, evolution can alter species ecological responses, and different aspects of evolution and ecology can interact to produce complex eco‐evolutionary dynamics under climate change. Here we review how evolution could alter ecological responses to climate change on species warm and cool range margins, where evolution could be especially important. We discuss different aspects of evolution in isolation, and then synthesize results to consider how multiple evolutionary processes might interact and affect conservation strategies. On species cool range margins, the evolution of dispersal could increase range expansion rates and allow species to adapt to novel conditions in their new range. However, low genetic variation and genetic drift in small range‐front populations could also slow or halt range expansions. Together, these eco‐evolutionary effects could cause a three‐step, stop‐and‐go expansion pattern for many species. On warm range margins, isolation among populations could maintain high genetic variation that facilitates evolution to novel climates and allows species to persist longer than expected without evolution. This ‘evolutionary extinction debt’ could then prevent other species from shifting their ranges. However, as climate change increases isolation among populations, increasing dispersal mortality could select for decreased dispersal and cause rapid range contractions. Some of these eco‐evolutionary dynamics could explain why many species are not responding to climate change as predicted. We conclude by suggesting that resurveying historical studies that measured trait frequencies, the strength of selection, or heritabilities could be an efficient way to increase our eco‐evolutionary knowledge in climate change biology.     

Botanical implications of bracken control

In addition to bracken, Pteridium aquilinum , and other ferns, only a restricted range of species among the natural flora suffer severe damage if sprayed with asulam in late summer. Of the plants associated with bracken in relatively well-drained situations those affected most by the herbicide include the three Ulex species, Rumex acetosella , young plants of Calluna vulgaris , some Compositae and grasses of the genera Poa, Holcus and Agrostis (though not A. setacea ). In damp and wet upland habitats, Cirsium palustre, Juncus effusus, Glyceria fluitans and a few other species are moderately susceptible to asulam but the majority of plants appear to be resistant or only slightly damaged.
Areas from which bracken has largely been removed tend to be invaded by such species as Deschampsia flexuosa, Holcus mollis, Chamaenerion augustifolium, Digitalis purpurea and Urtica dioica . If grazing pressure is low the absence of competition enables woody species to regenerate from seed. Colonization and subsequent spread of the perennial plants is influenced by what was present as ground flora before the bracken was killed and by the depth of bracken litter.
Asulam is considered to be a useful aid to the management of nature reserves where encroachment by bracken needs to be checked. Its most widespread use for bracken control is likely to be on upland farms to clear bracken from marginal land with the aim of increasing and improving pasture. Adjoining woodland and moorland habitats may suffer from resulting changes in land management. Moreover, indiscriminate aerial spraying of asulam over large areas may affect local populations of plants, notably ferns, in the more open areas not protected by a canopy of bracken.     

Local range boundaries vs. large‐scale trade‐offs: climatic and competitive constraints on tree growth

Species often respond to human‐caused climate change by shifting where they occur on the landscape. To anticipate these shifts, we need to understand the forces that determine where species currently occur. We tested whether a long‐hypothesised trade‐off between climate and competitive constraints explains where tree species grow on mountain slopes. Using tree rings, we reconstructed growth sensitivity to climate and competition in range centre and range margin tree populations in three climatically distinct regions. We found that climate often constrains growth at environmentally harsh elevational range boundaries, and that climatic and competitive constraints trade‐off at large spatial scales. However, there was less evidence that competition consistently constrained growth at benign elevational range boundaries; thus, local‐scale climate‐competition trade‐offs were infrequent. Our work underscores the difficulty of predicting local‐scale range dynamics, but suggests that the constraints on tree performance at a large‐scale (e.g. latitudinal) may be predicted from ecological theory.