Phenology of temperate trees in tropical climates

Several North American broad-leaved tree species range from the northern United States at 47°N to moist tropical montane forests in Mexico and Central America at 15–20°N. Along this gradient the average minimum temperatures of the coldest month (T _Jan), which characterize annual variation in temperature, increase from –10 to 12°C and tree phenology changes from deciduous to leaf-exchanging or evergreen in the southern range with a year-long growing season. Between 30 and 45°N, the time of bud break is highly correlated with T _Jan and bud break can be reliably predicted for the week in which mean minimum temperature rises to 7°C. Temperature-dependent deciduous phenology—and hence the validity of temperature-driven phenology models—terminates in southern North America near 30°N, where T _Jan>7°C enables growth of tropical trees and cultivation of frost-sensitive citrus fruits. In tropical climates most temperate broad-leaved species exchange old for new leaves within a few weeks in January-February, i.e., their phenology becomes similar to that of tropical leaf-exchanging species. Leaf buds of the southern ecotypes of these temperate species are therefore not winter-dormant and have no chilling requirement. As in many tropical trees, bud break of Celtis, Quercus and Fagus growing in warm climates is induced in early spring by increasing daylength. In tropical climates vegetative phenology is determined mainly by leaf longevity, seasonal variation in water stress and day length. As water stress during the dry season varies widely with soil water storage, climate-driven models cannot predict tree phenology in the tropics and tropical tree phenology does not constitute a useful indicator of global warming.     

Global patterns of insect herbivory in gap and understorey environments,and their implications for woody plant carbon storage

Insect herbivory is thought to favour carbon allocation to storage in juveniles of shade‐tolerant trees. This argument assumes that insect herbivory in the understorey is sufficiently intense as to select for storage; however, understoreys might be less attractive to insect herbivores than canopy gaps, because of low resource availability and – at temperate latitudes – low temperatures. Although empirical studies show that shade‐tolerant species in tropical forests do allocate more photosynthate to storage than their light‐demanding associates, the same pattern has not been consistently observed in temperate forests. Does this reflect a latitudinal trend in the relative activity of insect herbivory in gap versus understorey environments? To date there has been no global review of the effect of light environment on insect herbivory in forests. We postulated that if temperature is the primary factor limiting insect herbivory, the effect of gaps on rates of insect herbivory should be more evident in temperate than in tropical forests; due to low growing season temperatures in the oceanic temperate forests of the Southern Hemisphere, the effect of gaps on insect herbivory rates should in turn be stronger there than in the more continental temperate climates of the Northern Hemisphere. We examined global patterns of insect herbivory in gaps versus understories through meta‐analysis of 87 conspecific comparisons of leaf damage in contrasting light environments. Overall, insect herbivory in gaps was significantly higher than in the understorey; insect herbivory was 50% higher in gaps than in understoreys of tropical forests but did not differ significantly between gaps and understories in temperate forests of either hemisphere. Results are consistent with the idea that low resource availability – and not temperature – limits insect herbivore activity in forest understoreys, especially in the tropics, and suggest the selective influence of insect herbivory on late‐successional tree species may have been over‐estimated.     

Ontogenetic and temporal variations in herbivory and defense of Handroanthus spongiosus (Bignoniaceae) in a Brazilian tropical dry forest

We compared the richness and abundance of free-feeding herbivore insects (sap-sucking and leaf-chewing), leaf herbivory damage, leaf toughness and total phenolic content between two ontogenetic stages (juvenile and reproductive) of Handroanthus spongiosus (Rizzini) S. O. Grose (Bignoniaceae) throughout the rainy season in a Brazilian seasonally dry tropical forest. Twenty marked individuals of H. spongiosus were sampled per ontogenetic stage in each period of the rainy season (beginning, middle, and end). Herbivore richness and abundance did not differ between ontogenetic stages, but higher percentage of leaf damage, higher concentration of phenolic compounds, and lower leaf toughness were observed for juvenile individuals. The greatest morphospecies abundance was found at the beginning of the rainy season, but folivory increment was higher at the end, despite the fact that leaf toughness and total phenolic content increased in the same period. No significant relationships between leaf damage and both total phenolic content and leaf toughness were observed. These results suggest that insect richness and abundance do not track changes in foliage quality throughout plant ontogeny, but their decrease along rainy season confirms what was predicted for tropical dry forests. The general trends described in the current study corroborate those described in the literature about herbivores and plant ontogeny. However, the lack of relationship between herbivore damage and the two plant attributes considered here indicates that the analyses of multiple defensive traits (the defense syndrome) must be more enlightening to determine the mechanisms driving temporal and spatial patterns of herbivore attack.     

Leaf phenology is associated with soil water availability and xylem traits in a tropical dry forest

In tropical dry forests, spatial heterogeneity in soil water availability is thought to determine interspecific differences in key components of resource use strategies, such as leaf phenology and xylem function. To understand the environmental drivers of variation in leaf phenology and xylem function, we explored the relation of soil water potential to topographic metrics derived from a digital elevation model. Subsequently, we compared nine xylem hydraulic, mechanical and storage traits in 18 species in three phenological classes (readily deciduous, tardily deciduous, and evergreen) in the dry tropical forest of Chamela, Mexico. Soil water potential was negatively correlated with elevation, insolation and water flow accumulation. Evergreen species characterized low-elevation moist sites, whereas deciduous species dominated hills and dry sites. Overall, evergreen species had lower xylem specific conductivity than deciduous species, and tardily deciduous species were different from readily deciduous and evergreen species in five of eight xylem traits. In dry tropical forests, water availability promotes divergence in leaf phenology and xylem traits, ranging from low wood density, evergreen species in moist sites to a combination of low wood density, readily deciduous species plus high wood density, tardily deciduous species in dry sites.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Litterfall seasonality and adaptive strategies of tropical and subtropical evergreen forests in China

中国热带和亚热带常绿林凋落物季节特征及适应策略 本研究收集了来自中国热带/亚热带常绿林共85个站点的凋落物量季节性变化数据,并采用线性回归、结构方程模型构建以及相位差分析等方法,综合探究中国热带/亚热带地区常绿阔叶林和针叶林叶片脱落对土壤水分、饱和水气压差和辐射强度等气候因子的响应机制。研究结果显示,在雨热同期和雨热异期两种热带/亚热带气候类型中,呈现出两种典型凋落物的物候类型(单峰季节型/双峰季节型)。在雨热同期气候条件下,光照强度和降水呈现季节性正相关,单峰的凋落物峰值和双峰的第一个峰值约出现在3–4月,不断增加的光照能促进新叶的萌发,老叶被代谢更强的新叶所替代,该类型属于一种最大程度利用光照来实现树木生长的自适应策略。双峰的第二个峰值出现在雨季末期,约在8–10月,是由不断增强的水分亏缺所导致的(常绿阔叶林：大气水分亏缺;常绿针叶林：土壤水分亏缺),这种类型是一种凋落老叶减少水分丢失来应对水分胁迫的自适应策略。在雨热异期气候条件下,光照强度和降水呈现季节性负相关,饱和水气压差与光照强度表现出一致的季节性动态变化,诱导了常绿阔叶林单峰和双峰物候的第一个凋落峰(约在3–4月),是一种权衡大气干旱胁迫和最大程度利用光照进行生长的综合自适应策略。在雨季初期,显著的土壤水分亏缺加速叶片凋落,诱导了常绿阔叶林双峰物候的第二个凋落峰(约在11月),属于凋落老叶应对土壤水分胁迫的自适应策略。这些研究结果可以为地球系统模式中热带物候的精确模拟提供重要参考。     

西双版纳热带季节雨林林窗内幼树叶生长与虫食动态

 研究了西双版纳热带季节雨林6种乔木幼树在林窗中的叶生长与叶虫食动态。6种幼树叶生长主要在雨季(5～10月),在旱季雾凉期(11～2月)叶停止生长。阳性树种旱季干热期(3～4月)开始叶生长,顶极树种至5月雨季初才开始叶生长。表明雾凉期低温抑制两类树种叶生长,干热期水分不足抑制顶极树种叶生长。叶生长同步性(每两个月的叶生长量变异系数C．V．)顺序为：望天树(Shorea chinensis,1.42)>金钩花(Pseuduvaria indochinensis,1.41)>八宝树(Duabanga grandiflora, 1.02)>云南石梓(Gmelina arborea,0.98)>团花树(Anthocephalus chinensis,0.84)和铁刀木(Cassia siamea,0.84)。旱季造成一些种类幼树出现落叶高峰,严重叶虫食也导致云南石梓在7～8月出现落叶高峰。叶虫食主要出现在雨季,团花树和云南石梓在干热期叶生长能减少昆虫取食,但此时铁刀木叶同步生长却不能降低食叶昆虫危害。6种幼树叶生长量年进程与叶虫食量年进程间存在正相关,其中望天树、金钩花、八宝树和团花树分别达到显著(p<0.05)或极显著(p<0.01)水平。望天树和金钩花在雨季初叶同步生长能减轻叶虫食。食叶昆虫偏爱取食幼叶,6种幼树平均幼叶的虫食量占总虫食量的72.9％,幼叶虫食速率平均为成熟叶的4.3倍。     

Increasing day-length induces spring flushing of tropical dry forest trees in the absence of rain

In many conspecific trees of >50 species highly synchronous bud break with low inter-annual variation was observed during the late dry season, around the spring equinox, in semideciduous tropical forests of Argentina, Costa Rica, Java and Thailand and in tropical savannas of Central Brazil. Bud break was 6 months out of phase between the northern and southern hemispheres and started about 1 month earlier in the subtropics than at lower latitudes. These observations indicate that "spring flushing", i.e., synchronous bud break around the spring equinox and weeks before the first rains of the wet season, is induced by an increase in photoperiod of 30 min or less. Spring flushing is common in semideciduous forests characterized by a 4-6 month dry season and annual rainfall of 800-1,500 mm, but rare in neotropical forests with a shorter dry season or lower annual precipitation. Establishment of new foliage shortly before the wet growing season is likely to optimize photosynthetic gain in tropical forests with a relatively short growing season.     

Do seasonal changes in light availability influence the inverse leafing phenology of the neotropical dry forest understory shrub Bonellia nervosa (Theophrastaceae)?

In tropical dry forests most plants are deciduous during the dry season and flush leaves with the onset of the rains. In Costa Rica, the only species displaying the opposite pattern is Bonellia nervosa. To determine if seasonal changes in light availability are associated with the leaf and reproductive phenology of this species, we monitored leaf production, survival, and life span, as well as flower and fruit production from April 2000 to October 2001 in Santa Rosa National Park. Leaf flushing and flower bud production took place shortly after the autumnal equinox when day length starts to decrease. Leaves began expansion at the end of the wet season, and plants reached 70 % of their maximum leaf area at the beginning of the dry season, maintaining their foliage throughout the entire dry period. Leaf shedding occurred gradually during the first three months of the wet season. Leaf flushing and shedding showed high synchrony, with leaf numbers being related to light availability. Maximum leaf production coincided with peaks in radiation during the middle of the dry season. Decreasing day length induces highly synchronous flower bud emergence in dry forest species, but this is the first study indicating induction of leaf flushing by declining day length.     

Leaf phenology in relation to canopy closure in southern Appalachian trees

Leaf phenology varies markedly across tree species of temperate deciduous forests. Early leafing in spring may increase light capture and carbon gain prior to canopy closure, allowing saplings to survive in understory sites deeply shaded in midsummer. We quantified sapling leaf phenology for 18 tree species and seasonal variation in understory light availability at three sites along a ridge-slope-cove landform gradient in the Great Smoky Mountains National Park. Early leafing species (e.g., Aesculus flava, Carpinus caroliniana) broke bud an average of 24 d before late leafers (e.g., Magnolia fraseri, Nyssa sylvatica). Canopy closure occurred 14-18 d earlier and summer understory light was on average 63-74% lower on intermediate and mesic sites than on the xeric site. Early leafing species intercepted 45-80% of their growing season photon flux before canopy closure vs. 8-15% for late leafers. However, earlier leafing increased exposure to freezing temperatures by 5.5% per week near the mean time of bud break. Early leafing is strongly correlated with midsummer shade, risk of freezing temperatures, and distribution on mesic sites across a "main spectrum" of 15 deciduous species. Differences in leaf phenology and resultant impacts on spring carbon gain may help determine tree shade tolerance and distribution in southern Appalachian forests.     

Phenology, Lignotubers, and Water Relations of Cochlospermum vitifolium, a Pioneer Tropical Dry Forest Tree in Costa Rica

We examined structural and physiological traits relevant to the phenology of the tropical dry forest (TDF) pioneer tree Cochlospermum vitifolium . Despite marked seasonality in rainfall, meristem activity occurred throughout the year. Leaves were produced almost continuously during the rainy season, while leaf shedding started early during drought, before changes in soil water content were observed. Phenological activity under drought included flowering and fruiting of leafless trees; bud break and shoot extension took place before the end of the dry season. Low wood density of C. vitifolium stems (0.17 g/cm³) and lignotubers (0.14 g/cm³) provided water and starch storage needed to support phenological events such as branch extension, leaf flushing, and reproduction during the dry season, and probably also contributed to survival following mechanical damage and fire, typical of early TDF successional stages. Lignotuber water and starch contents showed substantial seasonal variation, declining from the beginning of the dry season to their lowest levels at the time of reproduction and dry-season flushing. Stems progressively replaced lignotubers as main storage organs as tree size increased. Evidence for a role of water stores in buffering daily water deficits was weak. Leaf water potentials remained above −1.2 MPa and stomatal conductance below 350 mmol/m²/s, suggesting that gas exchange during the rainy season was limited to prevent xylem cavitation. Leaf shedding occurred when early-morning and mid-day Ψ_L converged at the rainy–dry season transition, without changes in lignotuber or soil water content, suggesting that leaves of C. vitifolium are closely tuned to atmospheric drought.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .     

Abundance and Leaf Damage Caused by Gall‐Inducing Insects in a Mexican Tropical Dry Forest1

We evaluated the temporal and spatial patterns of abundance and the amount of damage caused by gall‐inducing insects (GII) in deciduous and riparian habitats in a seasonal tropical dry forest in Mexico. Plants occurring in these habitats differ in their phenology and moisture availability. Deciduous habitats are seasonal and xeric, while riparian habitats are aseasonal and mesic. We found 37 GII species and each one was associated with a specific plant species. In total, 19 species (51.3%) were present in deciduous habitats, 13 species (35.2%) in riparian habitats, and only 5 species (13.5%) occurred in both. Abundance and leaf damage by GII were greater in deciduous than in riparian habitats during the wet season. For each GII species that occurred in both habitats, host plant species supported greater abundance and leaf damage by GII in deciduous habitats during the wet season. These results indicate a greater association of GII species with host plants in deciduous than in riparian habitats during the wet season. In riparian habitats, 11 plant species (61.1%) had greater density of GII in the dry than in the wet season. Similarly, leaf damage by GII was significantly greater in the dry than in the wet season in riparian habitats for 12 plant species (66.7%). Dry forest plants of riparian habitats presented two peaks of leaf‐flushing: GII colonized leaves produced in the first peak at the beginning of the wet season, and accumulated or recolonized leaves in the second peak at the beginning of the dry season. The levels of leaf damage by GII detected in this study in the rainy season were considerably higher than those obtained for folivorous insects in other neotropical forests, suggesting that this GII guild might have an important impact on their host plant species in this tropical community.     

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.     

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.     

Patterns of leaf-pathogen infection in the understory of a Mexican rain forest: incidence, spatiotemporal variation, and mechanisms of infection

This study assessed the levels of damage by leaf pathogens and their variability in terms of host species, space (four mature forest sites) and season of the year (dry and rainy), and the mechanisms of infection in the understory of the Los Tuxtlas tropical rain forest. Sixty-five percent of the species surveyed in the dry season (N = 49) and 64.9% of those surveyed in the rainy season (N = 57) were damaged by fungi. Leaf area damaged per plant, on average, was <1% (range: 0.25-20.52%). There was considerable variation in the degree of infection among species, but not among sites and seasons. The survey showed that 43% of the leaves were damaged by herbivores and pathogens concurrently, 16% showed damage by insect herbivory alone, and only 1.4% of the sampled leaves showed damage by pathogens alone. Pathogenicity assays experimentally confirmed that the predominant mechanism of fungal establishment was wounding, such as that caused by herbivory (or other similar sources), and only rarely did infection occur through direct contact (without wounds). The results revealed the omnipresence of leaf fungal infection, although with low damage per plant, and the importance of herbivorous insects in the facilitation of fungal infection in tropical understory plants.     

Phenology of Tree Species in Bolivian Dry Forests

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub–canopy forest tree species were recorded monthly over a two–year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub–canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub–canopy tree species produced larger animal– or gravity–dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerio received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry forests.     

Evidence for facultative deciduousness in Colophospermum mopane in semi‐arid African savannas

Leaf phenology dictates the time available for carbon assimilation, transpiration and nutrient uptake in plants. Understanding the environmental cues that control phenology is therefore vital for predicting climate‐related changes to plant and ecosystem function. In contrast to temperate systems, and to a lesser degree, tropical forest systems, the cues initiating leaf drop in tropical savannas are poorly studied. We investigated the cues for leaf fall in a tropical monodominant arid savanna species, Colophospermum mopane, using an irrigation experiment. We tracked soil moisture, solar radiation, air temperature, leaf water status, leaf health and leaf carbon balance through the dry season in both irrigated and control plants. Water was the primary cue driving leaf loss of C. mopane rather than temperature or light. Trees watered throughout the dry season retained their canopies. These leaves remained functional and continued photosynthesis throughout the dry season. Leaf carbon acquisition rates did not decline with leaf age but were affected by soil moisture availability and temperature. Leaf loss did not occur when leaf carbon gain was zero, or when a particular leaf carbon threshold was reached. Colophospermum mopane is facultatively deciduous as water availability determines leaf drop in this widespread arid savanna species. Obligate deciduosity is not the only successful strategy in climates with a long dry season.     

Reproductive biology of Hechtia schottii, a dioecious Bromeliaceae, in Mexico

Hechtia schottii is a terrestrial, rosetofilous, dioecious, polycarpic succulent herb, that grows mainly in shrubby associations, and less frequently, in secondary low caducifolious forests, both on calcareous soils or limestone outcrops in Yucatan and Campeche States, Mexico. We studied phenology, floral and pollination biology, and breeding system at Calcehtok, Yucatan, during two flowering seasons. Plants bloom mainly during the dry season (November-April) and disperse seeds during the rainy season (May-October). Both floral morphs have diurnal anthesis; pollen is removed ca. 1 h after anthesis starts and both floral morphs are visited by several insect species, especially bees, but results suggest that the introduced honey bee, Apis mellifera, is the pollinator. Controlled crossings show that the species is functionally dioecious and requires to be serviced by pollinators based on fruit setting only in unassisted cross pollination crosses.     

Seasonal development of secondary xylem and phloem in <Emphasis Type="Italic">Schizolobium parahyba</Emphasis> (Vell.) Blake (Leguminosae: Caesalpinioideae)

The cambial activity and periodicity of secondary xylem and phloem formation have been less studied in tropical tree species than in temperate ones. This paper describes the relationship between seasonal cambial activity, xylem and phloem development, and phenology in Schizolobium parahyba, a fast growing semideciduous seasonal forest tree from southeastern Brazil. From 2002 to 2003, wood samples were collected periodically and phenology and climate were recorded monthly in the same period. S. parahyba forms annual growth increments in wood, delimited by narrow initial parenchyma bands. The reduction of the cambial activity to a minimum correlates to the dry season and leaf fall. The higher cambial activity correlates to the wet season and the presence of mature leaves. In phloem, a larger conductive region was observed in the wet season, when the trees were in full foliage. The secondary phloem did not exhibit any incremental zone marker; however, we found that the axial parenchyma tends to form irregular bands.     

Fruiting and flushing phenology in Asian tropical and temperate forests: implications for primate ecology

In order to understand the ecological adaptations of primates to survive in temperate forests, we need to know the general patterns of plant phenology in temperate and tropical forests. Comparative analyses have been employed to investigate general trends in the seasonality and abundance of fruit and young leaves in tropical and temperate forests. Previous studies have shown that (1) fruit fall biomass in temperate forest is lower than in tropical forest, (2) non-fleshy species, in particular acorns, comprise the majority of the fruit biomass in temperate forest, (3) the duration of the fruiting season is shorter in temperate forest, and (4) the fruiting peak occurs in autumn in most temperate forests. Through our comparative analyses of the fruiting and flushing phenology between Asian temperate and tropical forests, we revealed that (1) fruiting is more annually periodic (the pattern in one year is similar to that seen in the next year) in temperate forest in terms of the number of fruiting species or trees, (2) there is no consistent difference in interannual variations in fruiting between temperate and tropical forests, although some oak-dominated temperate forests exhibit extremely large interannual variations in fruiting, (3) the timing of the flushing peak is predictable (in spring and early summer), and (4) the duration of the flushing season is shorter. The flushing season in temperate forests (17–28 % of that in tropical forests) was quite limited, even compared to the fruiting season (68 %). These results imply that temperate primates need to survive a long period of scarcity of young leaves and fruits, but the timing is predictable. Therefore, a dependence on low-quality foods, such as mature leaves, buds, bark, and lichens, would be indispensable for temperate primates. Due to the high predictability of the timing of fruiting and flushing in temperate forests, fat accumulation during the fruit-abundant period and fat metabolization during the subsequent fruit-scarce period can be an effective strategy to survive the lean period (winter).