A novel framework for disentangling the scale‐dependent influences of abiotic factors on alpine treeline position

Low‐temperature growth limitation largely determines alpine treeline position globally, but treeline elevation also varies locally at a range of scales in response to multiple biotic and abiotic factors. In this study, we conceptualise how variability in treeline elevation is related to abiotic factors that act as thermal modifiers, physiological stressors, or disturbance agents. We then present a novel analytical framework for quantifying how abiotic factors influence treeline elevation at different spatial scales using New Zealand Nothofagus treelines as a case study. We delineated Nothofagus treelines in a GIS, along which we extracted data for treeline elevation and eight abiotic explanatory variables at 54 000 points. Each location was classified at each of five spatial scales based on nested river catchments, ranging from large regional to small hillslope catchments. We used hierarchical linear models to partition the variation in both treeline elevation and the eight abiotic variables by spatial scale, and then quantified the relationships between these at each spatial scale in turn. Nothofagus treeline elevation varied from 800–1740 m a.s.l. across New Zealand. Abiotic factors explained 82% of the variation in treeline elevation at the largest (regional) scale and 44–52% of variation at the four finer scales. Broad‐scale variation in Nothofagus treeline elevation was strongly associated with thermal modifiers, consistent with the idea that treelines coincide with a temperature‐driven, physiological limit. However, much of the finer‐scale variation in treeline elevation was explained by a combination of thermal, physiological stress‐related, and disturbance variables operating at different spatial scales. The conceptual model and analytical methods developed here provide a general framework for understanding treeline variation at different spatial scales.     

A re-assessment of high elevation treeline positions and their explanation

In this review I first compile data for the worldwide position of climate-driven alpine treelines. Causes for treeline formation are then discussed with a global perspective. Available evidence suggests a combination of a general thermal boundary for tree growth, with regionally variable “modulatory” forces, including the presence of certain taxa. Much of the explanatory evidence found in the literature relates to these modulatory aspects at regional scales, whereas no good explanations emerged for the more fundamental global pattern related to temperature per se, on which this review is focused. I hypothesize that the life form “tree” is limited at treeline altitudes by the potential investment, rather than production, of assimilates (growth as such, rather than photosynthesis or the carbon balance, being limited). In shoots coupled to a cold atmosphere, meristem activity is suggested to be limited for much of the time, especially at night. By reducing soil heat flux during the growing season the forest canopy negatively affects root zone temperature. The lower threshold temperature for tissue growth and development appears to be higher than 3°C and lower than 10°C, possibly in the 5.5–7.5°C range, most commonly associated with seasonal means of air temperature at treeline positions. The physiological and developmental mechanisms responsible have yet to be analyzed. Root zone temperature, though largely unknown, is likely to be most critical. Received: 3 October 1997 / Accepted: 14 April 1998     

Sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change at landscape and local scales

The sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change are increasingly discussed in terms of climate change, often forgetting that climate is only one aspect of environmental variation. As treeline heterogeneity increases from global to regional and smaller scales, assessment of treeline sensitivity at the landscape and local scales requires a more complex approach than at the global scale. The time scale (short‐, medium‐, long‐term) also plays an important role when considering treeline sensitivity. The sensitivity of the treeline to a changing environment varies among different types of treeline. Treelines controlled mainly by orographic influences are not very susceptible to the effects of warming climates. Greatest sensitivity can be expected in anthropogenic treelines after the cessation of human activity. However, tree invasion into former forested areas above the anthropogenic forest limit is controlled by site conditions, and in particular, by microclimates and soils. Apart from changes in tree physiognomy, the spontaneous advance of young growth of forest‐forming tree species into present treeless areas within the treeline ecotone and beyond the tree limit is considered to be the best indicator of treeline sensitivity to environmental change. The sensitivity of climatic treelines to climate warming varies both in the local and regional topographical conditions. Furthermore, treeline history and its after‐effects also play an important role. The sensitivity of treelines to changes in given factors (e.g. winter snow pack, soil moisture, temperature, evaporation, etc.) may vary among areas with differing climatic characteristics. In general, forest will not advance in a closed front but will follow sites that became more favourable to tree establishment under the changed climatic conditions.     

Treeline dynamics in relation to climatic variability in the central Tianshan Mountains, northwestern China

Aim Climate variability may be an important mediating agent of ecosystem dynamics in cold, arid regions such as the central Tianshan Mountains, north‐western China. Tree‐ring chronologies and the age structure of a Schrenk spruce (Picea schrenkiana) forest were developed to examine treeline dynamics in recent decades in relation to climatic variability. Of particular interest was whether tree‐ring growth and population recruitment patterns responded similarly to climate warming. Location The study was conducted in eight stands that ranged from 2500 m to 2750 m a.s.l. near the treeline in the Tianchi Nature Reserve (43°45′?43°59′ N, 88°00′?88°20′ E) in the central Xinjiang Uygur Autonomous Region, northwestern China. Methods Tree‐ring cores were collected and used to develop tree‐ring chronologies. The age of sampled trees was determined from basal cores sampled as close as possible to the ground. Population age structure and recruitment information were obtained using an age–d.b.h. (diameter at breast height) regression from the sampled cores and the d.b.h. measured on all trees in the plots. Ring‐width chronologies and tree age structure were both used to investigate the relationship between treeline dynamics and climate change. Results Comparisons with the climatic records showed that both the radial growth of trees and tree recruitment were influenced positively by temperature and precipitation in the cold high treeline zone, but the patterns of their responses differed. The annual variation in tree rings could be explained largely by the average monthly minimum temperatures during February and August of the current year and by the monthly precipitation of the previous August and January, which had a significant and positive effect on tree radial growth. P. schrenkiana recruitment was influenced mainly by consecutive years of high minimum summer temperatures and high precipitation during spring. Over the last several decades, the treeline did not show an obvious upward shift and new recruitment was rare. Some trees had established at the treeline at least 200 years ago. Recruitment increased until the early 20th century (1910s) but then decreased with poor recruitment over the past several decades (1950–2000). Main conclusions There were strong associations between climatic change and ring‐width patterns, and with recruitments in Schrenk spruce. Average minimum temperatures in February and August, and total precipitation in the previous August and January, had a positive effect on tree‐ring width, and several consecutive years of high minimum summer temperature and spring precipitation was a main factor favouring the establishment of P. schrenkiana following germination within the treeline ecotone. Both dendroclimatology and recruitment analysis were useful and compatible to understand and reconstruct treeline dynamics in the central Tianshan Mountains.     

Regional and local effects of disturbance and climate on altitudinal treelines in northern Patagonia

Abstract. In this field study we analysed the regional and local scale effects of disturbance and climate on altitudinal treelines dominated by Nothofagus pumilio in northern Patagonia. We compared two regions west and east of the Andes at 40° S, slopes with warm vs cool aspects and undisturbed vs locally disturbed treelines. This spatial framework allowed us to test (1) for differences among treelines affected by different types of local disturbance and (2) the traditional hypothesis that low temperature limits treeline. Contingency tables and ANOVA showed that local disturbance occurred more frequently than expected on slopes with cool aspects, steep slope angles and concave slope configuration. Disturbed treelines were locally lowered with longer ecotones and lower krummholz growth rates and vegetation cover than undisturbed treelines. Three‐way ANOVA showed the significant influences of study area (regional climate) and aspect (local climate) on treeline elevation, krummholz growth rates and density, tree density and vegetation cover, while accounting for local disturbance. Treeline elevations were higher east of the Andes reflecting the more continental climate in Argentina than in Chile, plus regional impacts of volcanic eruptions. Tree density and vegetation cover were greater west of the Andes reflecting greater precipitation in Chile. Within study areas, local climate had different influences on treeline elevations and krummholz growth rates west and east of the Andes. We predict that increased tree growth and upslope advance of treeline in response to global warming is more likely in Chile than in Argentina near 40° S, unless precipitation increases east of the Andes. To test these predictions, we recommend research be stratified to account for the influences of local disturbance, which may confound climatic impacts. In northern Patagonia, suitable control (undisturbed) study sites will most likely be found at upper slope positions with low slope angles, simple microtopography and straight topographic configuration.     

Gradient analysis of reversed treelines and grasslands of the Valles Caldera,New Mexico

Objective: : Treeless meadows and parks are widespread but poorly understood features of the montane vegetation of the western USA. These communities frequently form reversed treelines where grassy valleys occur below forested slopes above. Our purpose was to assess the environmental correlates of such treelines, as well as patterns in the composition and diversity of grasslands and forest margins in the Valles Caldera National Preserve. Location: Valles Caldera National Preserve (35°50′‐36°00’ N, 106°24′‐106°37’ W, 2175–3150 m), Jemez Mountains, New Mexico, USA. Methods: We conducted a gradient analysis based on 200 nested quadrats on transects crossing reversed treelines and spanning the compositional heterogeneity of grasslands. We used cluster analysis and non‐metric multidimensional scaling to assess relationships between compositional variation and environmental variables. Results: We found strong, highly significant relationships of the vegetation to gradients in slope inclination, soil texture, moisture, nutrient availability, and nighttime minimum temperatures. Reversed treelines are most strongly associated with shifts in the thermal regime, exhibit weaker relationships with soil texture and nutrient content, and show no relationship with gravimetric soil moisture. Gradients in aspect, soil moisture, and annual mean temperature are associated with compositional variation within grasslands and forest margins. Conclusions: Lower nightly minimum temperatures and fewer consecutive frost‐free days resulting from cold‐air drainage may prevent tree seedling establishment in valley bottoms via photo‐inhibition, tissue damage, or frost heaving. Fine‐textured soils may also impede tree seedling establishment in valley bottoms. These findings lay the groundwork for experimental and physiological tests of these potential causes of these reversed treelines.     

Temperature responses of carbon mineralization in conifer forest soils from different regional climates incubated under standard laboratory conditions

¹⁴C‐labelled straw was mixed with soils collected from seven coniferous forests located on a climatic gradient in Western Europe ranging from boreal to Mediterranean conditions. The soils were incubated in the laboratory at 4°, 10°, 16°, 23° and 30 °C with constant moisture over 550 days. The temperature coefficient (Q₁₀) for straw carbon mineralization decreased with increasing incubation temperatures. This was a characteristic of all the soils with a difference of two Q₁₀ units between the 4–10° and the 23? 30 °C temperature ranges. It was also found that the magnitude of the temperature response function was related to the period of soil incubation. Initial temperature responses of microbial communities were different to those shown after a long period of laboratory incubation and may have reflected shifts in microbial species composition in response to changes in the temperature regime. The rapid exhaustion of the labile fractions of the decomposing material at higher temperatures could also lead to underestimation of the temperature sensitivity of soils unless estimated for carbon pools of similar qualities. Finally, the thermal optima for the organic soil horizons (Of and Oh) were lower than 30 °C even after 550 days of incubation. It was concluded that these responses could not be attributed to microbial physiological adaptations, but rather to the rates at which recalcitrant microbial secondary products were formed at higher temperatures. The implication of these variable temperature responses of soil materials is discussed in relation to modelling potential effects of global warming.     

Notes on the Effect of Cutting Bracken (Pteris aquilina L.)

Background: Undisturbed high elevation treelines follow a common growing season isotherm, irrespective of latitude. Small stature plants thrive at much higher elevations because they grow in a favourable microclimate near the ground, whereas trees are aerodynamically coupled to free atmospheric circulation, hence the uniform treeline elevation in a given region.

Aims: I argue that the treeline results from tree architecture and not from a tree-specific inferior physiology. At tissue level, all cold adapted higher plants (including winter crops) face similar temperature related limitations and are constrained by similar thermal thresholds.

Methods: In order to explore this hypothesis, winter rape and winter oat were grown in the field and the daily rate of leaf expansion was measured during early winter, with temperatures similar to those at treeline at the beginning of the growing season (monthly mean temperature of 5 °C).

Results: Leaves of these winter crops only grew when the degree hours above 5 °C for a given day exceeded zero. Rape and oats showed very similar responses. The data support a common temperature threshold for tissue formation in winter crops and treeline trees.

Conclusions: A literature survey revealed lower temperature thresholds (close to freezing point) in arctic and Antarctic algae. It appears that the formation of complex, partly lignified tissue does not happen below 5 °C, whereas such temperatures exert little constraints on photosynthesis. The study illustrates that tissue growth in winter crops can help understand growth of treeline trees, given that both exhibit similar minimum temperature requirements for meristem functioning.     

TEMPERATURE RESPONSES AND EVOLUTION OF THERMAL TRAITS IN CLADOPHOROPSIS MEMBRANACEA (SIPHONOCLADALES,CHLOROPHYTA)1

Temperature tolerances and relative growth rates were determined for different isolates of the tropical to warm temperate seaweed species Cladophoropsis membranacea (C. Agardh) Boergesen (Siphonodadales, Chlorophyta) and some related taxa. Most isolates of C membranacea survived undamaged at 18° C for at least 8 weeks. Lower temperatures (5°–15°C) were tolerated for shorter periods of time but caused damage to cells. All isolates survived temperatures up to 34° C, whereas isolates from the eastern Mediterranean and Red Sea survived higher temperatures up to 36°C. Growth occurred between 18° and 32° C, but an isolate from the Red Sea had an extended growth range, reaching its maximum at 35°C. Struvea anastomosans (Harvey) Piccone & Grunow, Cladophoropsis sundanensis Reinbold, and an isolate of C. membranacea from Hawaii were slightly less cold- tolerant, with damage occurring at 18°C. Upper survival temperatures were between 32° and 36° C in these taxa. Temperature response data were mapped onto a phylogenetic tree. Tolerance for low temperatures appears to be a derived character state that supports the hypothesis that C. membranacea originated from a strictly tropical ancestor. Isolates from the Canary Islands, which is near the northern limit of distribution, are ill adapted to local temperature regimes. Isolates from the eastern Mediterranean and Red Sea show some adaptation to local temperature stress. They are isolated from those in the eastern Atlantic by a thermal barrier at the entrance of the Mediterranean.     

Thermal adaptations to extreme freeze–thaw cycles in the high tropical Andes

Temperature plays a key role in the biology of ectotherms, including anurans, which are found at higher elevations in the tropics than anywhere in the temperate zone. High elevation tropical environments are characterized by extreme daily thermal fluctuation including high daily maxima and nightly freezing. Our study investigated the contrasting operative temperatures of the anurans Telmatobius marmoratus and Pleurodema marmoratum in different environmental contexts at the same elevation and biome above 5,200 m. Telmatobius marmoratus avoids extremes of daily temperature fluctuation by utilizing thermally buffered aquatic habitat at all life stages, with minimal operative temperature variation (range: 4.6–8.0°C). Pleurodema marmoratum, in contrast, experienced operative temperatures from ?3.5 to 44°C and has one of the widest thermal breadths reported for any tropical frog, from >32°C (critical thermal maximum) to surviving freezing periods of 1 and 6 hr down to ?3.0°C. Our findings expand experimental evidence of frost tolerance in amphibians to the widespread Neotropical family Leptodactylidae, the first such evidence of frost tolerance in a tropical amphibian. Our study identifies three strategies (wide thermal tolerance breadth, use of buffered microhabitats, and behavioral thermoregulation), which allow these tropical frogs to withstand the current wide daily thermal fluctuation above 5,000 m.a.s.l. and which may help them adapt to future climatic changes. Abstract in Spanish is available with online material     

Limited prospects for future alpine treeline advance in the Canadian Rocky Mountains

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere     

Persistence and production of perennial grasses under water deficits and extreme temperatures: importance of intraspecific vs. interspecific variability

Extreme climatic events are expected to increase in frequency and magnitude as a consequence of global warming. Grasslands cover a large proportion of the European continent and contribute to both agricultural production and ecosystem services through inter and intraspecific genetic variability. This study analysed the effects of summer droughts and heat waves on the persistence and production of perennial forage grasses. Mediterranean and temperate populations of Dactylis glomerata L. and Festuca arundinacea (Schreb.) were compared at both Mediterranean and temperate sites in France. By manipulating canopy temperatures and water availability, grass swards in the field were subjected to cumulative summer and spring water deficits (CSSWD) ranging from 329 to 707 mm to test different projected climatic conditions and extreme summer events. Under controlled summer heat waves (6–21 days at a mean daily canopy temperature higher than 30–35 °C), there was no increase in membrane damage to surviving aerial tissues. Plant stress was thus mainly generated through greater soil water deficit. Under the greatest CSSWD, annual biomass production was reduced on average by 60% and 30% with temperate and Mediterranean populations, respectively. Thresholds for a significant increase in summer tiller mortality were seen at CSSWD higher than 450 mm for temperate populations and 550 mm for Mediterranean populations. The latter displayed lower predawn leaf water potentials in summer and recovered through intense tillering in the subsequent seasons. Under the most extreme CSSWD, fewer than 20% of tillers of temperate populations survived and their nitrogen uptake ability was drastically altered. The higher potential productivity of Mediterranean populations in winter was associated with greater frost sensitivity. The identification of thresholds for vulnerability and the determination of the role of genetic diversity will improve the management of plant resilience and the design of new plant material to cope with climate change.     

Thermal response in pre‐imaginal biology of Ochlerotatus albifasciatus from two different climatic regions

The biological processes on mosquito could be variable in response to local climatic characteristics. The thermal effects on time and the rate larval development, immature survival and adult size in local populations of Ochlerotatus albifasciatus (Macquart) (Diptera: Culicidae) from cold (Sarmiento) and temperate (Buenos Aires) regions from Argentina were evaluated. This species affects livestock production and human health. Larvae of both regions were placed in breeding thermal baths (11–32 °C range). Development and survival were recorded daily until adult emergence. The development temperature threshold and thermal constant for Sarmiento (4.59 ± 3.08 °C, 204.08 ± 7.83 degree days) was lower and higher than Buenos Aires, respectively (8.06 ± 1.81 °C, 149.25 ± 2.6 degree days). At cold temperatures (11–16 °C), Sarmiento larvae demonstrated 5 days faster development and higher survival (56%) than Buenos Aires (15%), whereas at warm temperatures (20–32 °C) were up to 2 days slower and similar survival (16% vs. 18%). The size did not show differences between populations. An Ochlerotatus albifasciatus population seems to present local thermal responses. The favourable temperature for survival and rate of development would vary within a cold or warm range, and these differential responses would explain the wide geographical distribution in different climatic regions of southern South America.     

Vegetation patterns,plant distribution and life forms across the alpine zone in southern Tierra del Fuego,Argentina

The alpine zone is examined at meso‐ and microscales in southern Tierra del Fuego (54°49′S), where the full zone is expressed. Mesoscale patterns were studied on opposing aspects, and microscale patterns were studied on a series of solifluction terraces, in a hanging valley overlooking the Beagle Channel. Plant cover and life form data were collected within 50‐m altitudinal bands on north and south aspects and comprehensive plant lists were compiled for each band. Topography and associated surface cover were recorded on the terraces. Six alpine plant communities, in lower and upper floristic zones, were differentiated with multivariate analyses and significantly related to five ecological factors. Equivalent communities were separated by approximately 185 m altitude on opposing aspects, which related to a soil temperature difference of approximately 3.0°C. The richness (and range) of 80 local vascular taxa (18.6% of the regional flora), decreased with increasing altitude (6.6 per 100 m); however, richness differed significantly with aspect (north: 5.6, south: 7.5). Upper altitudinal limits (approximately 1250 m a.s.l.), were associated with a midsummer isotherm of approximately 1.7°C. Chamaephytes and hemicryptophytes dominated throughout but the tall tussock form was conspicuously absent. Reasons for this are discussed in the context of the Nothofagus treeline, which conformed to a midsummer isotherm of only approximately 6.0°C. Such patterns are at variance with those found in the oceanic subantarctic islands, other oceanic perhumid temperate mountain regions and tropical high mountains. However, the microscale pattern of fines, pebbles, stones and rock across the active solifluction terraces, with dense vegetation on their steep risers, had a clear affinity with that of other subantarctic regions. Inferences that alpine systems of the Southern Hemisphere are necessarily equivalent to those at similar northern latitudes are cautioned against. Likewise, such comparisons within the Southern Hemisphere may also be invalid.     

Strong topographic sheltering effects lead to spatially complex treeline advance and increased forest density in a subtropical mountain region

Altitudinal treelines are typically temperature limited such that increasing temperatures linked to global climate change are causing upslope shifts of treelines worldwide. While such elevational increases are readily predicted based on shifting isotherms, at the regional level the realized response is often much more complex, with topography and local environmental conditions playing an important modifying role. Here, we used repeated aerial photographs in combination with forest inventory data to investigate changes in treeline position in the Central Mountain Range of Taiwan over the last 60 years. A highly spatially variable upslope advance of treeline was identified in which topography is a major driver of both treeline form and advance. The changes in treeline position that we observed occurred alongside substantial increases in forest density, and lead to a large increase in overall forest area. These changes will have a significant impact on carbon stocking in the high altitude zone, while the concomitant decrease in alpine grassland area is likely to have negative implications for alpine species. The complex and spatially variable changes that we report highlight the necessity for considering local factors such as topography when attempting to predict species distributional responses to warming climate.     

A temperate pollinator with high thermal tolerance is still susceptible to heat events predicted under future climate change

1. Mutualisms may be particularly vulnerable to climate change as interacting species are likely to respond differently, which could destabilise interactions. 2. Temperate zone insects typically experience mean temperatures below their thermal optima, making them less vulnerable than tropical insects to small increases in mean temperature. However, they are likely to experience a higher frequency of extreme heat events, putting mutualism persistence in jeopardy. 3. This study investigated the potential impacts of climate change on Pleistodontes imperialis, a temperate Australian fig wasp that pollinates Port Jackson figs (Ficus rubiginosa). Wasp emergence and longevity were measured at temperatures ranging from those commonly experienced in nature (25 °C) to high values (> 40 °C) that are currently infrequent, but which are becoming more common with climate change. 4. Wasp emergence was unaffected by temperatures up to 39 °C, but it declined drastically above 39 °C. Adult longevity was unaffected by temperatures up to 30 °C, but decreased at 35 °C and above. Low humidity reduced wasp longevity across all temperatures. 5. Fitness reductions were observed at temperatures ～5 °C above the summer daily mean maximum, suggesting that P. imperialis has a high thermal tolerance, but is vulnerable to extreme heat. Figs located in the shade may provide protected microhabitats under hot conditions. 6. Tropical pollinators may be threatened by small increases in mean temperature. In contrast, it is shown here that temperate pollinators may face a different primary threat from climate change – the increasing frequency of extreme heat events – despite their higher thermal tolerances.     

Assessing and modeling seed germination of Mediterranean wildflowers for low input landscape restoration

Native species are recommended for use in landscape restoration because they adapt well to the local pedo‐climatic conditions. Despite the high biodiversity in the Mediterranean, the use of native plants is hampered by the limited knowledge of their seed germination. This is particularly true for a number of plants which are appropriate for creating species‐rich herbaceous communities. In this study, seeds of 35 species were collected in different roadside and degraded sites in rural and urban areas. Two experiments were carried out to determine the influence of light and thermal conditions on seed germination. In the first experiment, seeds of 17 species were tested at different temperatures (5, 15, and 25°C). At 15°C, seed germination was tested under both dark and light conditions. In the second experiment, the germination of 30 species was tested under alternating temperatures (25/15°C) and dark/light conditions. The responses of the various species differed in relation to thermal levels and light conditions, e.g., Bartsia trixago did not germinate in the dark at constant temperatures (5, 15, and 25°C), while in the light (15°C) and at alternating temperatures (25/15°C) in light and dark conditions, germination was over 60%. In both experiments, Tragopogon porrifolius and Triticum ovatum showed the highest germination rate (≥88%). With the sole exception of Medicago orbicularis, all members of the Fabaceae showed no or low germination. The definition of the germination requirements of some Mediterranean species, highlighted in these experiments, provides useful information for the creation of low input green areas and environmental restoration using these species.     

Large-scale phytogeographical patterns in East Asia in relation to latitudinal and climatic gradients

Aim This paper aims at determining how different floristic elements (e.g. cosmopolitan, tropical, and temperate) change with latitude and major climate factors, and how latitude affects the floristic relationships between East Asia and the other parts of the world. Location East Asia from the Arctic to tropical regions, an area crossing over 50° of latitudes and covering the eastern part of China, Korea, Japan and the eastern part of Russia. Methods East Asia is divided into forty‐five geographical regions. Based on the similarity of their world‐wide distributional patterns, a total of 2808 indigenous genera of seed plants found in East Asia were grouped into fourteen geographical elements, belonging to three major categories (cosmopolitan, tropical and temperate). The 50°‐long latitudinal gradient of East Asia was divided into five latitudinal zones, each of c. 10°. Phytogeographical relationships of East Asia to latitude and climatic variables were examined based on the forty‐five regional floras. Results Among all geographical and climatic variables considered, latitude showed the strongest relationship to phytogeographical composition. Tropical genera (with pantropical, amphi‐Pacific tropical, palaeotropical, tropical Asia–tropical Australia, tropical Asia–tropical Africa and tropical Asia geographical elements combined) accounted for c. 80% of the total genera at latitude 20°N and for c. 0% at latitude 55–60°N. In contrast, temperate genera (including holarctic, eastern Asia–North America, temperate Eurasia, temperate Asia, Mediterranean, western Asia to central Asia, central Asia and eastern Asia geographical elements) accounted for 15.5% in the southernmost latitude and for 80% at 55–60°N, from where northward the percentage tended to level off. The proportion of cosmopolitan genera increased gradually with latitude from 5% at the southernmost latitude to 21% at 55–60°N, where it levelled off northward. In general, the genera present in a more northerly flora are a subset of the genera present in a more southerly flora. Main conclusions The large‐scale patterns of phytogeography in East Asia are strongly related to latitude, which covaries with several climatic variables such as temperature. Evolutionary processes such as the adaptation of plants to cold climates and current and past land connections are likely responsible for the observed latitudinal patterns.     

Treeline form – a potential key to understanding treeline dynamics

Aim Treelines occur globally within a narrow range of mean growing season temperatures, suggesting that low‐temperature growth limitation determines the position of the treeline. However, treelines also exhibit features that indicate that other mechanisms, such as biomass loss not resulting in mortality (dieback) and mortality, determine treeline position and dynamics. Debate regarding the mechanisms controlling treeline position and dynamics may be resolved by identifying the mechanisms controlling prominent treeline spatial patterns (or ‘form’) such as the spatial structure of the transition from closed forest to the tree limit. Recent treeline studies world‐wide have confirmed a close link between form and dynamics. Location The concepts presented refer to alpine treelines globally. Methods In this review, we describe how varying dominance of three general ‘first‐level’ mechanisms (tree performance: growth limitation, seedling mortality and dieback) result in different treeline forms, what ‘second‐level’ mechanisms (stresses, e.g. freezing damage, photoinhibition) may underlie these general mechanisms, and how they are modulated by interactions with neighbours (‘third‐level’ mechanisms). This hierarchy of mechanisms should facilitate discussions about treeline formation and dynamics. Results We distinguish four primary treeline forms: diffuse, abrupt, island and krummholz. Growth limitation is dominant only at the diffuse treeline, which is the form that has most frequently responded as expected to growing‐season warming, whereas the other forms are controlled by dieback and seedling mortality and are relatively unresponsive. Main conclusions Treeline form provides a means for explaining the current variability in treeline position and dynamics and for exploring the general mechanisms controlling the responses of treelines to climatic change. Form indicates the relative dependence of tree performance on various aspects of the external climate (especially summer warmth versus winter stressors) and on internal feedbacks, thus allowing inferences on the type as well as strength of climate‐change responses.     

Light,temperature, dry after‐ripening and salt stress effects on seed germination of Phleum sardoum (Hackel) Hackel

Phleum sardoum is an endemic psammophilous species of Sardinia, growing exclusively on coastal sandy dunes. The effect of glumes on seed germination, germination requirements at constant (5–25°C) and alternating (25/10°C) temperatures, both in the light (12/12 h) and in the dark were evaluated, as well as the effect of a dry after‐ripening period (90 days at 25°C), the salt stress effect (0–600 mmol NaCl) and its recovery on seed germination. The presence of glumes reduced final germination percentages. For fresh naked seeds, high germination percentages were observed at 10°C. Dry after‐ripening increased germination rate at low temperatures, but did not affect final germination percentages. NaCl determined a secondary salt‐induced dormancy which recovery interrupted only partially. Our results highlighted that this species has its optimum of germination during autumn–winter when, under a Mediterranean climate, water availability is highest and soil salinity levels are minimal.