Landscape patterns and environmental controls of deciduousness in forests of central Panama

Aim Dry season deciduousness affects intra‐ and inter‐annual patterns of carbon, water and energy balance in seasonal tropical forests. Because it is affected by rainfall, temperature and solar radiation, deciduousness may be an indicator of the response of vegetation to climate change. Better understanding of how spatial patterns of deciduousness are affected by climate and other environmental gradients will improve the ability to predict responses to climate change. This study develops remote sensing methods for quantifying tropical forest deciduousness and examines the relationship between deciduousness and environmental factors in semi‐deciduous tropical forest. Location Central Panama. Methods I applied spectral mixture analysis (SMA) and the normalized difference vegetation index (NDVI) to Landsat images to predict deciduousness which was ground‐truthed with field observations of the percentage of overstorey deciduous trees. Using predicted deciduousness from SMA, patterns of deciduousness at three spatial scales were analysed. I determined how deciduousness varied spatially with rainfall and geological substrate. Results Both SMA and NDVI had strong correlations (r > 0.9) with field observations of deciduousness. On a landscape scale, deciduousness increased as rainfall decreased, but geological substrate altered this relationship. On some geological substrates, deciduousness was much greater than expected for a given rainfall total or showed a slight but significant increase with rainfall. At an intermediate spatial scale, there were highly deciduous patches from 3 to 250 ha in size embedded in non‐deciduous forest, which may have resulted from topography, soil variation or past land use. Main conclusions Dry season deciduousness can be accurately quantified using satellite images indicating that remote sensing can be a valuable tool for detecting change and understanding ecosystem processes in tropical forests from landscape to regional scales.     

Latitudinal pattern of mountain vegetation zonation in southern and eastern Asia

A new scheme of altitudinal and latitudinal vegetation zonation is proposed for eastern Asia. The latitudinal patterns of mountain vegetation zonation show a clear boundary at ca. 20°–30° N. For the tropical mountains south of 20° N, the altitudinal series includes tropical lowland, tropical lower montane, and tropical upper montane zones. For the temperate mountains north of 30° N, the series includes temperate lowland, temperate lower montane, and temperate upper montane zones. The mountains located between 20° and 30° N show a transitional zonation pattern; the lower two zones are comparable to the lower two of the tropical zonation (tropical lowland and tropical lower montane), and the upper two zones are comparable to those of the temperate zonation (temperate lower montane and temperate upper montane). The tropical upper montane zone is not found north of 20°–30° N, while the tropical lower montane zone reaches down to sea level and constitutes the temperate lowland zone. Thus the zonation between 20° and 30° N includes tropical lowland, tropical lower montane/temperate lowland, temperate lower montane, and temperate upper montane zones. The latitudinal series of lowland rain forests follows the scheme of climatic division into tropical, subtropical/warm-temperate, cool-temperate and cold-temperate, with a shift of the respective life forms, evergreen, evergreen notophyllous, deciduous, and evergreen needle-leaved. The tropical lower montane forest can be correlated to the horizontal subtropical/ warm-temperate zone. The temperate altitudinal and latitudinal zonations above 30° N are correlated and show an inclined parallel pattern from high altitudes in the south to low altitudes down to sea level in the north.     

Deciduousness in tropical trees and its potential as indicator of climate change: A review

In tropical forests, deciduousness is an outcome of integrated effect of drought, tree characteristics and soil moisture conditions and thus it is a reliable indicator of seasonal drought experienced by different tree species. Variations in the deciduousness are associated with several ecophysiological characteristics, such as varying allocation pattern of metabolic products, resource capture and conservation, water relations and stem water storages, annual carbon sequestration, timing of reproductive event initiation, extent of separation of vegetative and reproductive events and leaf strategies, and it helps in maintenance of water balance and protection of tree organs during the seasonal drought. Tropical forests support mosaics of tree functional types showing marked differences in the duration of deciduousness (from leaf exchanging to >8 months deciduous), as a result of varying degree of water stress experienced by physiognomy, distribution and wood anatomy of tropical trees. Wide variations in deciduousness in the same species growing at different sites suggest the high sensitivity of tropical trees to small changes in growing habitat. In the present review we have explored the ecological significance of deciduousness in tropical trees with emphasis on: (a) inter- and intraspecies plasticity in deciduousness, (b) various capacity adaptations related with the duration of deciduousness, (c) relationship between tree stem water status and deciduousness, and (d) probable effect of impending climate change on tropical trees. An attempt has also been made to establish deciduousness as climate change indicator in the dry tropics. There is need to develop capabilities to detect and predict the impact of climate change on deciduousness through long-term phenological network in tropics. Remote sensing techniques can generate valuable ecological information such as leaf level drought response and phenological patterns. Deciduousness has the potential to emerge as an important focus for ecological research to address critical questions in global modeling, monitoring, and climate change.     

Variation in evergreen and deciduous species leaf phenology in Assam, India

In the present study phenological activities such as leaf and shoot growth, leaf pool size and leaf fall were observed for 3 years (March 2007–March 2010) in 19 tree species (13 evergreen and 6 deciduous species) in a wet tropical forest in Assam, India. The study area receives total annual average rainfall of 2,318 mm of which most rain fall (>70 %) occurs during June–September. Both the plant groups varied significantly on most of the shoot and leaf phenology parameters. In general, growth in deciduous species initiated before the evergreen species and showed a rapid shoot growth, leaf recruitment and leaf expansion compared to evergreen species. Leaf recruitment period was significantly different between evergreen (4.2 months) and deciduous species (6.8 months). Shoot elongation rate was also significantly different for evergreen and deciduous species (0.09 vs. 0.14 cm day^?1 shoot^?1). Leaf number per shoot was greater for deciduous species than for evergreen species (34 vs. 16 leaves). The average leaf life span of evergreen species (328 ± 32 days) was significantly greater than that of deciduous species (205 ± 16 days). The leaf fall in deciduous species was concentrated during the winter season (Nov–Feb), whereas evergreens retained their leaves until the next growing season. Although the climate of the study area supports evergreen forests, the strategies of the deciduous species such as faster leaf recruitment rate, longer leaf recruitment time, faster shoot elongation rate during favorable growing season and short leaf life span perhaps allows them to coexist with evergreen species that have the liberty to photosynthesize round the year. Variations in phenological strategies perhaps help to reduce the competition among evergreen and deciduous species for resources in these forests and enable the coexistence of both the groups.     

Quantifying the deciduousness of tropical forest canopies under varying climates

Abstract. Deciduousness is an important functional attribute of tropical trees, reflecting climatic conditions. Precisely quantifying and mapping deciduousness in tropical forests will be necessary for calibrating remote sensing images which attempt to assess canopy properties such as carbon cycling, productivity, or chlorophyll content. We thus set out to assess the degree of canopy deciduousness in three moist, semi‐deciduous tropical forests in central Panama. One site is a 6‐ha research plot near the Atlantic coast of Panama, where rainfall is 2830 mm/yr. The second site is a 50‐ha plot on Barro Colorado Island, near the center of the isthmus of Panama, where rainfall is 2570 mm/yr, and the final site is a 4‐ha plot near the Pacific coast of Panama, where rainfall is 2060 mm/yr. At each site, a random sample of trees from all canopy species (those with individuals ≥ 30 cm DBH) were visited and scored for deciduousness three times during the 1997 dry season. The estimated peak fraction of deciduous individuals in the canopy at the wetter site was 4.8%, at the intermediate site, 6.3%, and at the drier site, 24.3%. The estimated fraction of crown area deciduous peaked at 3.6%, 9.7%, and 19.1% at the wet, medium, and dry sites respectively. The percentage of canopy species that was deciduous –14%, 28%, and 41%–was much higher than the percentage of deciduous individuals, because not all individuals of deciduous species were deciduous. During the 1999 dry season, every individual of all the deciduous species was visited at the two drier sites, and the total number of deciduous trees observed closely matched the estimated numbers based on the smaller 1997 samples.     

Deciduous leaf drop reduces insect herbivory

Deciduous leaf fall is thought to be an adaptation that allows plants living in seasonal environments to reduce water loss and damage during unfavorable periods while increasing photosynthetic rates during favorable periods. Observations of natural variation in leaf shedding suggest that deciduous leaf fall may also allow plants to reduce herbivory. I tested this hypothesis by experimentally manipulating leaf retention for Quercus lobata and observing natural rates of herbivory. Quercus lobata is primarily deciduous although individuals show considerable natural variation in leaf retention. Oak saplings with no leaves through winter experienced reduced attack by cynipid gall makers the following spring. This pattern was consistent with the positive correlation between natural leaf persistence and gall numbers. These cynipids do not overwinter on the leaves that trees retain through winter, although they appear to use persistent leaves as oviposition cues. If these results are general for woody plants in continental temperate habitats, they suggest that an important and unrecognized consequence of deciduous leaf shedding may be a reduction in herbivore damage, and that this effect should be included in models of deciduous and evergreen behavior.     

Respirationsintensität in Stäinmen, Zweigen und Blältern von Laubbäumen im tropischen Regenwald und in temperierten Wäldern

Respiratory Activity in Stem, Branches and Leaves of Evergreen Trees in Tropical Rain Forest and of Deciduous Trees in Temperate Climate.— Reduced to the same temperature, the respiratory activity of different parts of tropical rain forest trees and of corresponding parts of temperate deciduous trees is approximately the same. In particular, the leaf-blades of Danish deciduous trees and of tropical rain forest trees from the lowland of Côte d'Ivoire (57° north. lat.) have nearly the same respiratory activity at 20°C: the blades of the extreme shade leaves about 0.1 mg CO₂, those of the extreme sun leaves about 0.8 mg CO₂ per 50 cm² (one side only) per hour at 20°C. This is in accordance with the fact that the blades of shade leaves, respectively Sim leaves, of the tropical rain forest trees are built in the same way as the blades of deciduous trees in our temperate climate. The respiratory activity shows that they have the same “concentration” of living cells, the same “concentration” of active plasma. The respiratory activity of trunk and branches depends mainly on three factors: diameter of the stem-section or branch-section, nutritional condition, and temperature. The respiratory activity of trunk and branches of hardwoods in temperate climate is also dependent on the season. The estimations on stems and branches from the temperature trees were made In July–August. There was in most cases a remarkable agreement in respiratory activity between temperate and tropical hardwoods: Branches with a diameter of about 0.5–2 cm have a respiration of about 70–100 mg CO₂ per kg fresh weight per hour at 20°C. Trunk sections with a diameter over 20 cm have a respiratory activity of about 3–5 mg CO₂, per kg fresh weight per hour at 20°C. In the older parts of a stem most of the cells are dead. The agreement in respiratory activity probably means that the “concentration” of living cells in the older parts of the stem of tropical and temperate hardwoods is the same.     

Periodicity of leaf growth and leaf dry mass changes in the evergreen and deciduous species of Southern Assam,India

The study described patterns of leaf dry mass change, leaf mass per area (LMA), relative growth rate and leaf life span (LL) for 14 evergreen and 7 deciduous species of a tropical forest of Southern Assam, India. Leaf expansion in both the groups was, in general, completed before June (i.e. well before the onset of monsoon rains). Although leaf dry mass during leaf initiation phase was significantly higher (P < 0.01) in evergreen species than in deciduous species, at the time of full leaf expansion, average leaf dry mass relative to the peak leaf dry mass, realised by the evergreen species was lower (66 %) than for deciduous species (76 %). Leaf dry mass increase in both groups continued after leaf full expansion. Evergreen species had a longer leaf dry mass steady phase than deciduous species (2–6 vs 2–3 months). Average LMA of mature leaves for evergreen species (77.43 g m^?2) was significantly greater than that of deciduous species (48.43 g m^?2). LL ranged from 165 days in Gmelina arborea (deciduous) to 509 days in Dipterocarpus turbinatus (evergreen). LMA was correlated positively with LL, indicating that evergreen species with higher leaf construction cost retain leaves for longer period to pay back. The average leaf dry mass loss before leaf shedding was greater (P < 0.01) for deciduous species (30.29 %) than for evergreen species (18.31 %). Although the cost of leaf construction in deciduous species was lower than for evergreen species, they replace leaves at a faster rate. Deciduous species perhaps compensate the cost involved in faster leaf replacement through higher reabsorption of dry mass during senescence, which they remobilise to initiate growth in the following spring when soil resources remain limiting.     

Energetics and water flux of the marbled velvet gecko (Oedura marmorata) in tropical and temperate habitats

The gecko Oedura marmorata was studied in two different climatic zones: the arid zone of central Australia and in the wet-dry tropics of northern Australia. Doubly labelled water was used to measure field metabolic rate (FMR) and water flux rates of animals in the field during the temperate seasons of spring, summer and winter, and during the tropical wet and dry seasons. FMRs were highest in the tropical wet season and lowest in the temperate winter. The geckos in central Australia expended less energy than predicted for a similarly sized iguanid lizard, but geckos from the tropics expended about the same amount of energy as predicted for an iguanid. Water flux rates of geckos from the arid zone were extremely low in all seasons compared to other reptiles, and although water flux was higher in tropical geckos, the rates were low compared to other tropical reptiles. The standard metabolic rates (SMRs) of geckos were similar between the two regions and among the seasons. Geckos selected higher body temperatures (T _bs) in a laboratory thermal gradient in the summer (33.5°C) and wet (33.8°C) seasons compared to the winter (31.7°C) and dry (31.4°C) seasons. The mean T _bs selected in the laboratory thermal gradient by geckos from the two regions were not different at a given time of year. The energy expended during each season was partitioned into components of resting metabolism, T _b and activity. Most of the energy expended by geckos from central Australia could be attributed to the effects of temperature on resting lizards in all three seasons, but the energy expended by tropical geckos includes a substantial component due to activity during both seasons. This study revealed variability in patterns of ecological energetics between populations of closely related geckos, differences which cannot be entirely attributed to seasonal or temperature effects. Received: 14 November 1997 / Accepted: 4 May 1998     

Using calls as an indicator for Antarctic blue whale occurrence and distribution across the southwest Pacific and southeast Indian Oceans

Understanding species distribution and behavior is essential for conservation programs of migratory species with recovering populations. The critically endangered Antarctic blue whale (Balaenoptera musculus intermedia) was heavily exploited during the whaling era. Because of their low numbers, highly migratory behavior, and occurrence in remote areas, their distribution and range are not fully understood, particularly in the southwest Pacific Ocean. This is the first Antarctic blue whale study covering the southwest Pacific Ocean region from temperate to tropical waters (32°S to 15°S). Passive acoustic data were recorded between 2010 and 2011 across the southwest Pacific (SWPO) and southeast Indian (SEIO) oceans. We detected Antarctic blue whale calls in previously undocumented SWPO locations off eastern Australia (32°S, 152°E) and within the Lau Basin (20°S, 176°W and 15°S, 173°W), and SEIO off northwest Australia (19°S, 115°E).In temperate waters, adjacent ocean basins had similar seasonal occurrence, in that calling Antarctic blue whales were present for long periods, almost year round in some areas. In northern tropical waters, calling whales were mostly present during the austral winter. Clarifying the occurrence and distribution of critically endangered species is fundamental for monitoring population recovery, marine protected area planning, and in mitigating anthropogenic threats.     

Novel psychrotolerant picocyanobacteria isolated from Chesapeake Bay in the winter

Picocyanobacteria are major primary producers in the ocean, especially in the tropical or subtropical oceans or during warm seasons. Many “warm” picocyanobacterial species have been isolated and characterized. However, picocyanobacteria in cold environments or cold seasons are much less studied. In general, little is known about the taxonomy and ecophysiology of picocyanobacteria living in the winter. In this study, 17 strains of picocyanobacteria were isolated from Chesapeake Bay, a temperate estuarine ecosystem, during the winter months. These winter isolates belong to five distinct phylogenetic lineages, and are distinct from the picocyanobacteria previously isolated from the warm seasons. The vast majority of the winter isolates were closely related to picocyanobacteria isolated from other cold environments like Arctic or subalpine waters. The winter picocyanobacterial isolates were able to maintain slow growth or prolonged dormancy at 4°C. Interestingly, the phycoerythrin‐rich strains outperformed the phycocyanin‐rich strains at cold temperature. In addition, winter picocyanobacteria changed their morphology when cultivated at 4°C. The close phylogenetic relationship between the winter picocyanobacteria and the picocyanobacteria living in high latitude cold regions indicates that low temperature locations select specific ecotypes of picocyanobacteria.     

Tree specific traits affect flowering time in Indian dry tropical forest

Dry tropical forest tree species show variations in leafless duration (i.e. deciduousness), stem wood density (SWD), leaf mass area (LMA) and leaf strategy index (LSI, reflecting resource use rate) to overcome water limitations. We investigated the role of these tree traits in the seasonal timing of flowering and subsequent fruiting. Flowering and fruiting time of 24 tree species was recorded over two consecutive annual cycles and their relationships with the abovementioned tree specific traits were examined across the species. In leaf-exchanging species having higher SWD and LMA, low LSI and short deciduousness, flowering coincides with leaf transitional state when vegetative growth is at its minimum, and fruit formation and leaf flushing are both supported at the same time. However, >4-months-deciduous species with lowest SWD and LMA, higher LSI and longer deciduousness showed predominantly dry season flowering, subsequent fruiting on leafless shoots and distinct separation of vegetative and flowering phenophases. In contrast, intermediate species (<2 months-deciduous, 2–4-months-deciduous) showed wider flowering range through summer, rainy, autumn or winter seasons. Fruiting duration varies considerably with variation in the flowering time; ca. 5–14 months in summer flowering species; 7–12 months in rainy flowering species; 6–10 months in autumn flowering species, 4–9 months in dry season flowering and 3–7 months in winter flowering species. In most species, fruit maturation occurred just prior to the onset of rains, ensuring seedling survival. The ability of tree species to withstand (leaf-exchange) or avoid (deciduousness) drought stress and varying seasonal flowering timings appear to be the principal mechanisms for successful survival and reproduction under extremely dry and seasonal climate. Since environmental characteristics affect flowering and fruiting either directly (e.g. through conditions in the habitat) or indirectly (e.g. through deciduousness, LMA, SWD and LSI), the impact of probable global climatic change will have long implications on reproduction of dry tropical trees.     

Aboveground biomass allocation patterns within Mediterranean sub-shrubs: A quantitative analysis of seasonal dimorphism

The monthly patterns of aboveground biomass allocation were studied in the branches of six Mediterranean sub-shrubs with different leaf phenology. Four of them were seasonally dimorphic species, and the remaining two were a winter deciduous and a cushion plant with photosynthetic stems. By the analysis of these species we aimed to identify different aboveground biomass allocation patterns within seasonally dimorphic species and to understand the role of seasonal dimorphism as a strategy to avoid the main stresses of mediterranean climate: summer drought and winter cold. The biomass allocation to the different living and photosynthetic fractions of 3-year-old branches was studied monthly for a minimum of 13 months per species. Leaf area (LA, mm²) and leaf mass per area (LMA, mg cm⁻²) measurements were used to characterize the diverse types of leaves of each species. Standing dead and senescent tissues accounted for a great percentage of the branch biomass of seasonally dimorphic species both during summer and winter. Different patterns of photosynthetic biomass allocation were found within the seasonally dimorphic species analysed. These patterns ranged from the moderate photosynthetic biomass oscillation of Salvia lavandulifolia to the almost deciduousness of Lepidium subulatum, and they were achieved by keeping alive, drying out or shedding different types of branches and leaves throughout the year. The formation of stress tolerant leaves and the reduction in the amount of photosynthetic biomass responded both to the occurrence of summer drought and winter cold. These results demonstrate that seasonal dimorphism is a flexible ecological strategy, as it comprises very different leaf phenologies and enables plants to escape both summer drought and winter cold.     

Network analysis of tree crowns distinguishes functional groups of Cerrado species

Deciduous, semideciduous and evergreen leaf phenological groups of Cerrado trees were studied using a representative network composed of nodes and links to uncover the structural traits of the crown. A node denotes the origin of a branch, and a link represents the branch emerging from a lateral bud. The network representation usually resulted in a graph with three links per node and twice as many links as nodes for each leaf phenological group. It was possible to identify four kinds of nodes according to the position and the number of links: initial, regular, emission and final nodes. The numbers of links and nodes and the distance between two kinds of nodes decreased from evergreen to deciduous species. A crown with a few nodes and links and a short distance between the kinds of nodes could facilitate the unfolding of foliage on leafless branches at the end of the dry season in deciduous trees. In contrast, foliage persistence in evergreens could facilitate the mass flow to new leaves produced during the entire year in a crown with a high number of links and nodes and with a large distance between nodes. There is a clear interdependence between the degree of leaf deciduousness and the crown structural traits in Cerrado tree species. Therefore, there are functional groups of trees in Cerrado vegetation that are characterized by a set of structural traits in the crown, which is associated with leaf deciduousness.     

Chilling and freezing stress in live oaks (Quercus section Virentes): intra- and inter-specific variation in PS II sensitivity corresponds to latitude of origin

Sensitivity to cold and freezing differs between populations within two species of live oaks (Quercus section Virentes Nixon) corresponding to the climates from which they originate. Two populations of Quercus virginiana (originating from North Carolina and north central Florida) and two populations of the sister species, Q. oleoides, (originating from Belize and Costa Rica) were grown under controlled climate regimes simulating tropical and temperate conditions. Three experiments were conducted in order to test for differentiation in cold and freezing tolerance between the two species and between the two populations within each species. In the first experiment, divergences in response to cold were tested for by examining photosystem II (PS II) photosynthetic yield (ΔF/F _m′) and non-photochemical quenching (NPQ) of plants in both growing conditions after short-term exposure to three temperatures (6, 15 and 30°C) under moderate light (400 μmol m⁻² s⁻¹). Without cold acclimation (tropical treatment), the North Carolina population showed the highest photosynthetic yield in response to chilling temperatures (6°C). Both ecotypes of both species showed maximum ΔF/F _m′ and minimum NPQ at their daytime growth temperatures (30°C and 15°C for the tropical and temperate treatments, respectively). Under the temperate treatment where plants were allowed to acclimate to cold, the Q. virginiana populations showed greater NPQ under chilling temperatures than Q. oleoides populations, suggesting enhanced mechanisms of photoprotective energy dissipation in the more temperate species. In the second and third experiments, inter- and intra-specific differentiation in response to freezing was tested for by examining dark-adapted F _v/F _m before and after overnight freezing cycles. Without cold acclimation, the extent of post-freezing declines in F _v/F _m were dependent on the minimum freezing temperature (0, −2, −5 or −10°C) for both populations in both species. The most marked declines in F _v/F _m occurred after freezing at −10°C, measured 24 h after freezing. These declines were continuous and irreversible over the time period. The North Carolina population, however, which represents the northern range limit of Q. virginiana, showed significantly less decline in F _v/F _m than the north central Florida population, which in turn showed a lower decline in Fv/F _m than the two Q. oleoides populations from Belize and Costa Rica. In contrast, after exposure to three months of chilling temperatures (temperate treatment), the two Q. virginiana populations showed no decline in F _v/F _m after freezing at −10°C, while the two Q. oleoides populations showed declines in F _v/F _m reaching 0.2 and 0.1 for Costa Rica and Belize, respectively. Under warm growth conditions, the two species showed different F ₀ dynamics directly after freezing. The two Q. oleoides populations showed an initial rise in F ₀ 30 min after freezing, followed by a subsequent decrease, while the Q. virginiana populations showed a continuous decrease in F ₀ after freezing. The North Carolina population of Q. virginiana showed a tendency toward deciduousness in response to winter temperatures, dropping 58% of its leaves over the three month winter period compared to only 6% in the tropical treatment. In contrast, the Florida population dropped 38% of its leaves during winter. The two populations of the tropical Q. oleoides showed no change in leaf drop during the 3-months winter (10% and 12%) relative to their leaf drop over the same timecourse in the tropical treatment. These results indicate important ecotypic differences in sensitivity to freezing and cold stress between the two populations of Q. virginiana as well as between the two species, corresponding to their climates of origin.     

Seasonal changes in energetics and torpor patterns in the subtropical blossom-bat Syconycteris australis (Megachiroptera)

Little is known about how animals from tropical and subtropical climates adjust their energy expenditure to cope with seasonal changes of climate and food availability. To provide such information, we studied the thermal physiology, torpor patterns and energetics of the nocturnal blossom-bat (Syconycteris australis 18 g) from a subtropical habitat in both summer and winter. In both seasons, S. australis frequently entered daily torpor at ambient temperatures between 12 and 25°C when food and water were withheld. Unlike patterns observed in temperate animals, mean minimum metabolic rates during torpor were lower in summer (0.47 ± 0.07 ml O₂ g⁻¹ h⁻¹) than in winter (0.75 ± 0.11 ml O₂ g⁻¹ h⁻¹). Body temperatures during torpor were regulated at 19.3 ± 1.0°C in summer and at 23.4 ± 2.0°C in winter. Torpor bout duration was significantly longer in summer (7.3 ± 0.6 h) than in winter (5.5 ± 0.3 h), but in both seasons, bout duration was not affected by ambient temperature. Consequently, average daily metabolic rates were also significantly lower in summer than in winter. Body temperatures and metabolic rates in normothermic bats did not change with season. Our findings on seasonal changes of torpor in this bat from the subtropics are opposite to those made for many species from cold climates which generally show deeper and longer torpor in winter and are often entirely homeothermic in summer. More pronounced torpor in subtropical S. australis in summer may be due to low or unpredictable nectar availability, short nights which limit the time available for foraging, and long days without access to food. Thus, the reversed seasonal response of this subtropical bat in comparison to temperate species may be an appropriate response to ecological constraints. Received: 6 May 1997 / Accepted: 19 October 1997     

Off-season uptake of nitrogen in temperate heath vegetation

In this field study we show that temperate coastal heath vegetation has a significant off-season uptake potential for nitrogen, both in the form of ammonium and as glycine, throughout winter. We injected ¹⁵N-ammonium and ¹⁵N 2×(¹³C)-glycine into the soil twice during winter and once at spring. The winter temperatures were similar to those of an average winter in the northern temperate region of Europe, with only few days of soil temperatures below zero or above 5°C. The vegetation, consisting of the evergreen dwarf shrub Calluna vulgaris, the deciduous dwarf shrub Salix arenaria, and the graminoids Carex arenaria and Deschampsia flexuosa, showed high root uptake of both forms of nitrogen, both 1 day after labelling and after a month, in species specific temporal patterns. Plant uptake of ¹³C was not significant, providing no further evidence of intact uptake of glycine. Translocation of the labelled nitrogen to shoots was generally evident after 1 month and increased as spring approached, with different translocation strategies in the three plant functional types. Furthermore, only the graminoids showed shoot growth during winter. Increasing plant nitrogen concentration from fall to spring at temperate heaths may, hence, be due to nitrogen uptake. Our results suggest that the potential for nitrogen uptake in plants at winter is of the same order of magnitude as at summer. Hence, winter nitrogen uptake in ecosystems in the temperate/boreal region should be considered when making annual nitrogen budgets of heath ecosystems, and the view of plant nutrient uptake as low in this climatic region during winter should be revised.     

Seasonality of peak metabolic rate in non‐migrant tropical birds

Birds that are year‐round residents of temperate and tropical regions have divergent life histories. Tropical birds have a slower ‘pace of life’, one characteristic of which includes lower peak metabolic rate and daily activity levels. Temperate resident birds are faced with seasonal variation in thermogenic demand. This challenge is met with seasonally increased peak metabolic rate during winter. These thermogenic demands are much lower in birds that are year‐round tropical residents. By measuring peak (summit) metabolic rate in tropical and temperate resident bird species during summer and winter, we asked whether tropical birds exhibit seasonality in peak metabolic rate, and if the direction of seasonality differs between tropical and temperate species. We measured summit metabolism in seven tropical and one temperate species during the winter and during the summer breeding season to test the hypothesis that summit metabolism of tropical residents would change seasonally. We consider whether metabolic seasonality is associated with breeding season for tropical species. We found that summit metabolism was significantly greater during the summer for most tropical residents, while the temperate resident matched several previous reports with higher summit metabolism in winter. We conclude that metabolic seasonality occurs in tropical residents and differs from temperate residents, suggesting that breeding during the summer may be driving relatively higher metabolism as compared to winter thermogenesis in temperate birds.     

Quantifying nationwide land cover and historical changes in forests of Nepal (1930–2014): implications on forest fragmentation

This study quantifies the nationwide land cover and long-term changes in forests and its implications on forest fragmentation in Nepal. The multi-source datasets were used to generate the forest cover information for 1930, 1975, 1985, 1995, 2005 and 2014. This study analyzes distribution of land cover, rate of deforestation, changes across forest types, forest canopy density and pattern of fragmentation. The land cover legend for 2014 is consisting of 21 classes: tropical dry deciduous sal forest, tropical moist deciduous sal forest, subtropical broad-leaved forest, subtropical pine forest, lower temperate broad leaved forest, upper temperate broad leaved forest, lower temperate mixed broad leaved forest, upper temperate mixed broad leaved forest, temperate needle leaved forest, subalpine forest, plantations, tropical scrub, subtropical scrub, temperate scrub, alpine scrub, grassland, agriculture, water bodies, barren land and settlements. The forest cover statistics for Nepal obtained in this study shows an area of 76,710 km² in 1930 which has decreased to 39,392 km² in 2014. A net loss of 37,318 km² (48.6%) was observed in last eight decades. Analysis of annual rate of net deforestation for the recent period indicates 0.01% during 2005–2014. An increase in the number of forest patches from 6925 (in 1930) to 42,961 (in 2014) was noticed. The significant observation is 75.5% of reduction in core 3 forest, whereas, patch, perforated and edge classes show the increase in percentage of fragmentation classes from 1930 to 2014. The results of this work will support the understanding of deforestation and its consequences on fragmentation for maintaining and improving the forest resources of Nepal.