Nutritional ecology of a browsing ruminant, the kudu (Tragelaphus strepsiceros), through the seasonal cycle

The activities and food selection of four hand-reared kudus were recorded in a large fenced enclosure containing natural savanna vegetation in the Nylsvley Nature Reserve, South Africa. Leaves of selected species were analysed chemically for crude protein, fibre constituents, phosphorus, condensed tannin and total polyphenols. Available protein and metabolizable energy were estimated allowing for potential antinutritional effects of tannins.
Leaves of palatable deciduous woody plants and herbaceous forbs formed the main dietary constituents during the late wet season. Foliage from palatable evergreens and robust forbs were added to the diet during the dry season. Towards the end of the dry season unpalatable species of evergreens were eaten. At the start of the growing season new leaves of otherwise unpalatable woody species formed the staple food source, together with fruits of Strychnos spp. Correspondingly, protein and digestible dry matter concentrations in the diet declined to reach a low at the end of the dry season.
Total daily food intake increased to compensate for reduced dietary quality during the dry season, until little edible foliage remained. While the estimated daily intake of protein remained well above maintenance requirements, the estimated metabolizable energy intake fell below requirements during the late dry season. Phosphorus intake may have been submaintenance in the dry season. Nutritional balance was dependent on the availability of particular vegetation components to serve as nutritional stepping stones during crucial times of the year. These included forbs during the late wet season, palatable evergreens in the dry season, and Strychnos fruits plus early-flushing woody plants during the dry season-wet season transition.     

Seasonal patterns of leaf production in co‐occurring trees with contrasting leaf phenology: time and quantitative divergences

Neotropical savannas (‘cerrados’) of Central Brazil are characterized by the coexistence of a large diversity of tree species with divergent phenological behaviors, which reflect a great diversity in growth strategies. In the present study time behavior and quantitative aspects of shoot growth, shoot mortality, and leaf longevity and production were analyzed in 12 woody species of contrasting leaf phenology, adopting a functional group approach where 12 species were categorized into three functional groups: evergreen, decidous and brevideciduous, according to their leaf phenology. Shoot growth and leaf production were seasonal for the three functional groups, differing in their time of occurrence, but being concentrated during the last months of the dry season. Shoot growth differed between evergreens and deciduous, as well leaf production. Evergreens had higher rates of shoot growth, produced a higher number of leaves and had longer leaf longevity (around 500 days against 300 days in deciduous and brevideciduous). Leaf longevity was associated with patterns of leaf production when accounting for all phenological groups studied. It was possible to identify different patterns of aerial growth in savanna phenological groups, providing evidence of great functional variability amongst the groups studied.     

Relationships between leaf deciduousness and flowering traits of woody species in the Brazilian neotropical savanna

The relationships between foliage permanence and flowering throughout the year were analyzed in 92 woody species of Cerrado vegetation categorized as either deciduous (DE), semideciduous (SD) or evergreen (EV). Flowering of DE, SD and EV species was investigated via three variables, measured over the course of the year: flowering duration (FLD), calculated as the number of months in flower in each species; flowering distribution (FDI), calculated as the number of species in flower per month; and flowering peak (FPE), defined as the four consecutive months yielding the highest number of species in flower. The months with the highest numbers of species in flower were October (52 species), September (50) and August (49). These months correspond to the period of transition from the dry season to the wet season. In the majority of species studied, seasonal climatic factors were strong enough to induce fruit formation in the dry season and seed dispersal in the following wet season, when sufficient water was available to support germination and plantlet growth. However, significant differences in FLD, FDI and FPE were found among the leaf phenological groups. High FLD in EV species is likely favored by the continuous input of resources from the year-round foliage. In contrast, DE species employ reserves of carbon, water and nutrients to form new leaves and flowers on a crown free of foliage at the end of the dry season. In DE species, their low FLD may reduce the impact of flowering on reserve consumption. SD species showed an intermediate level of foliage persistence, resulting in intermediate FLD values. In addition, SD species exhibited a different pattern of flowering distribution from those of DE and EV species. Many SD species have two flowering periods per year. The first period occurs when the crowns are full of leaves, in the middle of the dry season in June, similar to EV species. The second occurs when only half of the original foliage area is present, near the peak of the dry season in September, similar to DE species. Therefore, despite a strong influence of seasonal climatic conditions on the flowering behavior of DE, SD and EV woody species of Cerrado vegetation, these leaf phenological groups differ significantly in FLD, FDI and FPE.     

Do the phenology and functional stem attributes of woody species allow for the identification of functional groups in the semiarid region of Brazil?

The phenology of tree species in environments that are subject to strong climatic seasonality is mainly determined by water availability, which may vary as a function of wood density. The relationship among phenology, water potential, wood density and the capacity of water storage in the stem were determined for woody species of caatinga vegetation (dry forest) in the semiarid region of NE Brazil. Leaf flush and fall, flowering and fruiting events were recorded over a 31-month period, and the water potential was measured over a two-year period. These data were related to precipitation, water availability in the soil and photoperiod. Seven deciduous species exhibited low wood density (DLWD,?<0.5?g?cm^?3), high capacity of water storage in the stem (until 250?% of the dry weight) and high water potential during the year, as opposed to 15 deciduous species that showed high wood density (DHWD,?≥0.5?g?cm^?3). Leaf flush, flowering and the fruiting of DHWD species were related to precipitation, whereas these phenological events occurred at the end of the dry season and/or the beginning of the rainy season for DLWD species and were related to the photoperiod. The two evergreen species showed variations of water potential that were intermediate between those of DHWD and DLWD deciduous species, leaf flush during the dry season and flowering at the end of dry season. These results suggest the existence of three functional groups: evergreen species, DHWD deciduous species and DLWD deciduous species.     

Modification of Vegetative Phenology in a Tropical Semi‐deciduous Forest by Abnormal Drought and Rain1

The control of vegetative phenology in tropical trees is not well understood. In dry forest trees, leaf abscission may be enhanced by advanced leaf age, increasing water stress, or declining photoperiod. Normally, it is impossible to dissect the effects of each of these variables because most leaves are shed during the early dry season when day length is near its minimum and leaves are relatively old. The 1997 El‐Niño Southern Oscillation caused a ten‐week long, severe abnormal drought from June to August in the semi‐deciduous forests of Guanacaste, Costa Rica. We monitored the effect of this drought on phenology and water status of trees with young leaves and compared modifications of phenology in trees of different functional types with the pattern observed during the regular dry season. Although deciduous trees at dry sites were severely water stressed (Ψstem < ‐7MPa) and their mesic leaves remained wilted for more than two months, these and all other trees retained all leaves during the abnormal drought. Many trees exchanged leaves three to four months earlier than normal during the wet period after the abnormal drought and shed leaves again during the regular dry season. Irrigation and an exceptional 70 mm rainfall during the mid‐dry season 1998/1999 caused bud break and flushing in all leafless trees except dormant stem succulents. The complex interactions between leaf age and water stress, the principal determinants of leaf abscission, were found to vary widely among trees of different functional types.     

Ecological and ecophysiological patterns in a pre-altiplano shrubland of the Andean Cordillera in northern Chile

We report on community structural, ecophysiological, phenological, and morphological measurements made on woody plant species in the high elevation pre-altiplano zone on the western slopes of the Andean Cordillera of northern Chile. Notwithstanding extreme conditions of low rainfall, high atmospheric vapour demand and diurnal temperature fluctuation, a diversity of habitats (associated with drainage and slope aspect), appreciable local plant species richness (28 woody perennial plant species in a small area), and an array of adaptive morphological ecophysiological and phenological traits are present among woody species in these shrublands. Family diversity was low with four families accounted for 82% of the species. A range of gas exchange and watering use efficiency strategies was present ranging from highly efficient CAM species with a carbon isotope discrimination (Δ) of 3.7–7.5‰ through C₃ species with varying stomatal and gas exchange characteristics with a Δ of 14.4 to 19.8‰. Drought-deciduous small-leaved amphistomatic species from arid slopes generally had high stomatal conductance and high carbon assimilation rates during the rainy season. These drought deciduous species were largely leafless and, with one exception had low water potentials, during the dry season. Wash and less xeric site species commonly had broader evergreen to semi-evergreen leaves, higher dry season water potentials, and relatively consistent and moderate rates of gas exchange throughout the year. For all species, intrinsic water use efficiency (as estimated from the inverse of c _i :c _a ratio) correlated positively with mean leaf width (broader leaves had a lower higher intrinsic WUE) and dry season water potential. The charismatic high altitude tree, Polylepis rugulosa (Roaceae), had a population structure that suggests highly episodic establishment of seedlings, likely in sequences of wet years. Little of the area of these significant shrublands is currently protected. It would be desirable to add areas of pre-altiplano shrublands to adjacent national parks to ensure the persistence of these important communities. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

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.     

Shoot growth phenology of co-existing evergreen and deciduous species in an oak forest

Shoot growth phenology was compared for the saplings of evergreen and deciduous woody species sharing the same microsite. Growth initiation occurred earlier in evergreens (among co-stratal species) while deciduous species completed their growth earlier. Shoot growth rate was significantly greater (P<0.01) for deciduous trees than evergreen trees. The amount of shoot elongations and shoot diameter was also significantly greater (P<0.01) for deciduous trees than evergreens. On the other hand, among shrubs the amount of shoot elongation and shoot diameter was greater for evergreens but the rate of elongation and diameter was more or less similar for both. The duration of shoot elongation and shoot diameter was significantly longer in evergreens than the deciduous species. Leaf packing (number of leaves per shoot) was significantly more dense in evergreen trees (P<0.01) than in deciduous tree species. Leaf packing was more dense in evergreen than deciduous shrubs but the difference was not significant. Leaf area (per individual leaf) at full expansion was significantly greater (P<0.01) in deciduous species. Leaf dry mass and specific leaf mass in the initial stage was significantly greater for evergreen species than for deciduous species. The number of buds/10 cm of shoot was higher in evergreens. However, the per cent mortality was also higher in them.     

Leaf growth pattern in evergreen and deciduous species of the Central Himalaya,India

Leaf growth patterns were investigated in 11 evergreen (with leaf life-spans of just more than 1 year) and 15 deciduous species, occurring along an elevational gradient of 600–2200 m elevation in the Central Himalaya. Records were made of the leaf initiation period, leaf population dynamics, leaf expansion, leaf mass changes, leaf longevity and related parameters. Species of both groups produced leaves at similar rates during March to April, the driest period of the year. Species of both groups had approximately fully developed foliage during the warm, wet period (mid-June to mid-September) of the monsoon. However, significant differences were found at group level in other characters: shoot length (19.5 cm per shoot for deciduous and 11.7 cm for evergreen species); leaf population per 10 cm shoot length (4.7 vs 15.0); leaf area (107.9 vs 41.4 cm²/ leaf); specific leaf mass (106.9 vs 191.3 g/m²); and leaf mass loss after the monsoon period, being rapid and higher (31.6%) in deciduous species and slow and limited in the evergreens (26.2%). However, species of the two groups showed considerable overlaps in the values of above characters. The evergreen species of the Central Himalaya resembled the deciduous species of the region more than the multi-year leaves of clearly evergreen species. The evergreens bear leaves throughout the year, but like deciduous species bear the cost of annual replacement of old leaves by new leaves. They seem to outcompete deciduous species by producing annually a greater mass of leaves of low-carbon cost (per unit leaf mass), which is capable of conducting photosynthesis all year round. A situation of less marked contrast between favourable and nonfavourable periods, with respect to temperature, seems to favour the leaf characters of the evergreens.     

热带喀斯特森林常绿和落叶榕树的水力特征和水分关系与抗旱策略

以热带喀斯特地区的直脉榕(Ficus orthoneura)和豆果榕(F.pisocarpa)为实验材料,研究了常绿和落叶树木枝条和叶片的解剖结构特征、光合水分特征和耐旱性的差异,目的在于探讨不同生活型榕树适应干旱生境的策略.直脉榕和豆果榕的叶片都有两层栅栏组织、游离状的海绵组织和钟乳体等旱生结构,同时叶片角质层蒸腾速率(gmin)和气孔导度(gs)相对较低.但与落叶的豆果榕相比,常绿的直脉榕的枝条木质部失去50％传导率的水势(P50)和gmin更低,表现出更保守的水分利用策略.总体上,两种榕树都表现出了对喀斯特干旱生境的良好适应,但是它们的适应策略表现出一定的差别.豆果榕通过落叶度过旱期,而直脉榕在结构和功能上比豆果榕更耐旱.抗旱策略和水分利用策略的不同导致两种榕树的生态位分异,减少了彼此间的水分竞争,有利于它们在喀斯特生境中共存.     

Seasonality and life cycles of woody plant-feeding noctuid moths (Lepidoptera: Noctuidae) in Mediterranean habitats

Abstract.

• 1 The annual cycle of noctuid moths feeding as larvae on the foliage of eighteen woody plant species (belonging to ten different families) was studied at two southeastern Spanish Mediterranean forest habitats. Two questions were addressed in this study. (1) Does the rigorous (hot and dry) summer season characteristic of the Mediterranean-type climate impose a constraint on noctuid life histories? (2) Are there detectable differences in abundance or phenological patterns between the noctuid assemblages feeding on evergreen and winter-deciduous trees and shrubs?
• 2 Regardless of their leaf persistence habit, the majority of woody species studied had short shoot growth and leaf production periods in spring. Abundance of noctuid larvae peaked in May-June, shortly after host plants started to produce new leaves. Virtually no noctuid larvae were found on the foliage of the woody species studied at other times of year. Abundance and seasonal pattern of occurrence of larvae were similar on evergreen and deciduous food plants.
• 3 In contrast with the unimodal seasonal pattern exhibited by larvae, the abundance of adult moths (assessed by light trapping) showed two distinct peaks in early summer and early autumn, and a marked minimum in mid-summer.
• 4 Most woody plant-feeding noctuids recorded in this study (86% of species, about 95% of individuals) were univoltine. Univoltine species fell into one of two distinct life history categories, corresponding to pre- and post-summer flight periods. These two contrasting phenologies were associated with a bimodality in the duration of the pupal stage (‘short’ versus ‘long’), and were closely related to taxonomical affiliation at the subfamily level.
• 5 Within the ‘short’ pupal duration group, some species mate and oviposit shortly after emergence and pass the dry season in the egg stage, while others remain as potentially active, non-reproductive adults over most of the summer and mate by the end of that season. Species in the ‘long’ pupal duration group pass the summer in pupal or prepupal stage.
• 6 In the Mediterranean habitats studied, the adversity of the summer dry season seems to have led to a woody plant-feeding noctuid species assemblage almost entirely made up of univoltine species that concentrate their larval phases when suitable food (young leaves) is most abundant, and ‘avoid’ mid-summer as a flight time by emerging either shortly before or shortly after the adverse summer drought period. Year-round foliage availability afforded by dominant evergreen plants does not seem to have influenced the seasonal organization of noctuid cycles in any substantial way.

     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.     

Ecophysiological responses of two closely related Magnoliaceae genera to seasonal changes in subtropical China

Aims Plants use a variety of hydraulic strategies to adapt to seasonal drought that differ by species and environmental conditions. The early-diverging Magnoliaceae family includes two closely related genera with contrasting leaf habits, Yulania (deciduous) and Michelia (evergreen), which naturally inhabit temperate and tropical regions, respectively. Here, we evaluate the hydraulic strategy of species from both genera that have been ex situ conserved in a subtropical region to determine how they respond to the novel cool–dry season climatic pattern.Methods We measured ecophysiological traits in five Michelia and five Yulania species conserved in the South China Botanical Garden in both wet and dry season conditions and monitored the whole-year sap flow for four of these species.Important findings We found that Magnoliaceae species that have been ex situ conserved in a subtropical climate did not suffer from excessive water stress due to the mild drought conditions of the dry season and the ecophysiological adjustments the species made to avoid this stress, which differed by leaf habit. Specifically, deciduous species completely shed their leaves during the dry season, while evergreen species decreased their turgor loss points, dry mass based photosynthetic rates, stomatal conductance and specific leaf areas (SLAs) compared to wet season measurements. In comparing the two distinct leaf habits during the wet season, the leathery-leaved evergreen species had higher leaf hydraulic conductance and leaf to sapwood area ratios than the papery-leaved deciduous species, while the deciduous species had greater hydraulic conductivity calculated on both a stem and leaf area basis, dry mass based photosynthetic rates, leaf nutrients, SLAs and stomatal sizes than the evergreen species. Interestingly, species from both genera maintained similar sap flow in the wet season. Both photosynthetically active radiation and vapour pressure deficit affected the diurnal patterns of sap flow in the wet season, while only vapour pressure deficit played a dominant role in the dry season. This study reveals contrasting hydraulic strategies in Yulania and Michelia species under subtropical seasonal conditions, and suggests that these ecophysiological adjustments might be affected more by leaf habit than seasonality, thus reflecting the divergent evolution of the two closely related genera. Furthermore, we show that Magnoliaceae species that are ex situ conserved in a subtropical climate are hydraulically sound, a finding that will inform future conservation efforts of this ancient family under the threat of climatic change.     

Environmental constraints on phenology and internal nutrient cycling in the Mediterranean winter-deciduous shrub Amelanchier ovalis Medicus

The functional adjustments of winter-deciduous perennials to Mediterranean conditions have received little attention. The objectives of this study were: (i) to determine whether Amelanchier ovalis, a winter-deciduous shrub of Mediterranean and sub-Mediterranean regions, has nutritional and phenological traits in common with temperate zone deciduous trees and shrubs and (ii) to determine the constraints of Mediterranean environmental conditions on these traits. Over two years, phenology and nitrogen, and phosphorus concentrations were monitored monthly in the crown of A. ovalis. Leaf longevity, survival and nutrient resorption from senescing leaves were used to infer nutrient use efficiency and retention times of nutrients within the crown. In A. ovalis, bud burst was much earlier than in temperate deciduous trees and shrubs. Most vegetative and reproductive growth occurred in spring. Limited phenological development took place during the summer drought period. Unexpectedly, leaf shedding was very gradual, which might be related to water shortages in summer. Leaf longevity, nutrient resorption from senescing leaves, and maximum leaf nutrient concentrations indicated that nutrient retention times were short and nutrient use efficiency was low compared to that found in temperate deciduous plants and co-occurring Mediterranean evergreens. A. ovalis exhibited phenological development appropriate for a Mediterranean climate, although its limited ability to retain nutrients likely restricts the types of sites that it can occupy.     

云南元江干热河谷木本植物的物候

在中国西南干热河谷的典型地段——元江干热河谷,连续3年观测了32种木本植物的枝条生长、叶片动态、花期、果期和果实类型。这些植物的枝条生长方式可以分为连续生长、枝条枯死、陡长和间歇生长4个类型。其中连续生长型占优势,包括13种植物,它们的枝条在雨季连续不断伸长。9种植物雨季的枝条伸长与连续生长型的相似,但它们顶部的枝条在旱季末期出现枯死现象。6种植物属于陡长型,在2周内完成抽枝,且一年只抽一次枝。4种植物属于间歇生长型,枝条在雨季来临后伸长一段时间,然后生长停滞,过一段时间后再接着伸长。从叶片物候类型看,元江干热河谷植被以落叶植物占优势。落叶植物中冷凉旱季（11月～2月）落叶植物占优势（19种）,而干热旱季（3—4月）落叶植物很少（4种）。除红花柴（Indigofera pulchella）和狭叶山黄麻（Trema angustifolia）从雨季中期开始脱落叶片外,其它30种植物从雨季末期开始脱落叶片,落叶期至少延续3个月以上。常绿植物脱落近1／3～1／2的当年生叶片。共有6种植物能在旱季末期长出新叶。常绿植物的叶面积、单个枝条上的总叶面积和枝条承载（总叶面积／枝条长度）比落叶植物小。虽然一年四季都有不同植物开花和结果,但多数植物（29种,占观测树种的91％）的花期集中在旱季和雨季初期,而果实（种子）成熟期从雨季末期延续到旱季末期和下个雨季初期。果实多为核果。     

Functional groups of woody species in semi‐arid regions at low latitudes

In seasonally dry environments, woody species have different survival strategies. However, little is known about how environmental variables affect the phenology and water dynamics of these species. We aim to understand which variables initiate the vegetative phenophases of species in a tropical semiarid climate at 3°S latitude, where variation in photoperiod is minimal and rainfall is seasonal. We hypothesize that groups of species with similar vegetative phenologies, under similar conditions, are functionally similar in terms of water storage and use. We analyse the relationship between functional characteristics related to the acquisition and utilization of water, such as wood density, water storage capacity, water potential and vegetative phenology. The attributes were ordered by multidimensional scaling, and a multiple response permutation procedure was used to test consistency of the groups. Canonical correspondence analysis and Mantel tests were used to evaluate the phenophase response to environmental variables. We found four functional groups: (i) deciduous low wood density, which lose 75% of their leaves one month before the end of the rains; (ii) evergreen high wood density; (iii) early deciduous high wood density, which lose 75% of their leaves one month after the end of the rains; and (iv) late deciduous high wood density, which lose 75% of their leaves two months after the end of the rains. As expected, the vegetative phenodynamics of the deciduous high wood density group were mainly influenced by water availability. The evergreens did not show a correlation with rainfall. Only leaf shedding of the late deciduous, and the vegetative phenophases of the evergreens, responded to an increase in temperature and photoperiod. Bud‐break responded to increased photoperiod and soil humidity in the deciduous low wood density group. The foliar periodicity groups can be explained by the presence of species that differ mainly in their mechanisms of water acquisition and use.     

Seasonal Foliage Changes in the Eastern Amazon Basin Detected from Landsat Thematic Mapper Satellite Images1

Seasonal changes in tropical forests are difficult to measure from the ground, especially in areas of high species diversity and low phenological synchrony. Satellite images, which integrate individual tree canopies and cover a large spatial extent, facilitate tests for stand-level canopy phenology. Variability in near-infrated radiance (TM bands 4 and 5) of several distinct vegetation types was used to detect seasonal changes in a series of three Landsat Thematic Mapper (TM) images from the wet season to the dry season in Marabá, Brazil (eastern Amazon basin). Despite different atmospheric and instrumental conditions among the images, spectral changes were distinguishable. A phenological process (leaf aging, leaf drop, water stress) was determined from the spectral changes for each vegetation type. Changes in the spectral properties suggest that during the dry season, upland terra firme forest increased the rate of leaf exchange and some riparian vegetation was deciduous. Terra firme forest that had been altered by penetration of fires from nearby pastures increased in leaf biomass over a 14-month period. This study shows that a time series of images can provide information on temporal changes in primary vegetation and guide field studies to investigate seasonal changes that may not be detectable from the ground.