Tree seedling development in tropical dry abandoned pasture and secondary forest in Costa Rica

Development of planted seedlings of four canopy tree species in recently abandoned pastures (mown and unmown) and in ca. 40-yr old secondary dry forest in Guana-caste National Park, Costa Rica, was studied from July 1989–June 1992. The species were the light-demanding Cedrela odorata and Swietenia macrophylla, and the shade-tolerant Hymenaea courbaril and Manilkara chicle. Seedling mortality was high and primarily correlated with dry season drought. After the first dry season, M. chicle and C. odorata showed < 5 % survival in the pasture and 20 % in the forest, but after 3 yr survival had decreased to 3% in all plots. Survival of S. macrophylla was highest in the mown pasture (45 % after 3 yr) and lowest in the forest (10 %). For H. courbaril, survival was 40–55 % under all three conditions. Growth rates were low, with a height increment of < 15 cm/yr, possibly due to low precipitation in 1990 and 1991. All species grew taller in the pasture than in the forest. Differences in soil depth, texture and drainage appeared to contribute to variation in the results both within and between plots. Herbivory contributed to the lower survival of C. odorata and S. macrophylla in the forest.     

Floral traits and pollination systems in the Caatinga, a Brazilian tropical dry forest

BACKGROUND AND AIMS: Pollination is a critical stage in plant reproduction and thus in the maintenance and evolution of species and communities. The Caatinga is the fourth largest ecosystem in Brazil, but despite its great extent and its importance few studies providing ecological information are available, with a notable lack of work focusing on pollination biology. Here, general data are presented regarding the frequency of pollination systems within Caatinga communities, with the aim of characterizing patterns related to floral attributes in order to make possible comparisons with data for plant communities in other tropical areas, and to test ideas about the utility of syndromes. This paper also intends to provide a reference point for further studies on pollination ecology in this threatened ecosystem. METHODS: The floral traits and the pollination systems of 147 species were analysed in three areas of Caatinga vegetation in northeastern Brazil, and compared with world-wide studies focusing on the same subject. For each species, floral attributes were recorded as form, size, colour, rewards and pollination units. The species were grouped into 12 guilds according to the main pollinator vector. Analyses of the frequencies of the floral traits and pollination systems were undertaken. KEY RESULTS: Nectar and pollen were the most common floral resources and insect pollination was the most frequent, occurring in 69.9 % of the studied species. Of the entomophilous species, 61.7 % were considered to be melittophilous (43.1 % of the total). Vertebrate pollination occurred in 28.1 % of the species (ornithophily in 15.0 % and chiropterophily in 13.1 %), and anemophily was recorded in only 2.0 %. CONCLUSIONS: The results indicated that the pollination systems in Caatinga, despite climatic restrictions, are diversified, with a low percentage of generalist flowers, and similar to other tropical dry and wet forest communities, including those with high rainfall levels.     

Former land-use and tree species affect nitrogen oxide emissions from a tropical dry forest

Species composition in successional dry forests in the tropics varies widely, but the effect of this variation on biogeochemical processes is not well known. We examined fluxes of N oxides (nitrous and nitric oxide), soil N cycling, and litter chemistry (C/N ratio) in four successional dry forests on similar soils in western Puerto Rico with differing species compositions and land-use histories. Forests patch-cut for charcoal 60 years ago had few legumes, high litter C/N ratios, low soil nitrate and low N oxide fluxes. In contrast, successional forests from pastures abandoned several decades ago had high legume densities, low litter C/N ratios, high mean soil nitrate concentrations and high N oxide fluxes. These post-pasture forests were dominated by the naturalized legume Leuceana leucocephala, which was likely responsible for the rapid N cycling in those forests. We conclude that agriculturally induced successional pathways leading to dominance by a legume serve as a mechanism for increasing N oxide emissions from tropical regions. As expected for dry regions, nitric oxide dominated total N oxide emissions. Nitric oxide emissions increased with increasing soil moisture up to about 30% water-filled pore space then stabilized, while nitrous oxide emissions, albeit low, continued to increase with increasing soil wetness. Inorganic N pools and net N mineralization were greatest during peak rainfalls and at the post-agricultural site with the highest fluxes. Soil nitrate and the nitrate/ammonium ratio correlated positively with average N oxide fluxes. N oxide fluxes were negatively and exponentially related to litter C/N ratio for these dry forests and the relationship was upheld with the addition of data from seven wet forests in northeastern Puerto Rico. This finding suggests that species determination of litter C/N ratio may partly determine N oxide fluxes across widely differing tropical environments.     

Relationships among time, frequency, and duration of flowering in tropical rain forest trees

Flowering patterns are defined by the timing, duration, and frequency of flowering. Plants, particularly in the tropics, vary enormously with respect to these main variables of flowering. We used data from 302 tree species in a wet tropical forest to test a series of predictions regarding timing, duration, and frequency of flowering and examined the effect of each variable on the other two. Because timing, duration, and frequency of flowering can be constrained by phylogeny, we analyzed the data before and after considering phylogenetic effects at the level of family. Flowering activity peaked in the first wet season from May to July, refuting our prediction of peak flowering during the dry season. Our prediction that most species should flower several times a year was supported when species flowering more or less continually throughout the year were included in this category. Our prediction that supra-annually flowering species should be the least frequent was also supported with some qualifications. As we predicted, species flowering several times a year bloomed relatively briefly per flowering episode. Our prediction of shorter flowering duration for species flowering in the dry season and for those with a temporal separation between flowering and vegetative growth was also supported. Furthermore, supra-annually flowering species flowered for a shorter duration than annually flowering species and had a higher probability of flowering in the dry season compared to episodically or annually flowering species. Phylogeny significantly constrained variation in flowering frequency, but not in flowering time or duration, among confamilial species.     

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.     

Growth of tree seedlings in a tropical dry forest in relation to soil moisture and leaf traits

"AimsThe growth of plant species in tropical dry forest (TDF) is expected to be largely governed by the availability of soil moisture. In this study we attempt to identify mechanisms by which seedlings of dry tropical trees cope with water stress by adjusting their leaf characteristics to water availability and micro environments, and address following questions: How are leaf traits and relative growth rate (RGR) of the dominant seedling species of TDF affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional traits with each other? Can leaf traits singly or in combination predict the growth rate of seedling species of TDF? The study was conducted in situ on four sites (viz., Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India. Methods Five leaf traits viz., specific leaf area (SLA), leaf dry matter content (LDMC), concentrations of leaf nitrogen (leaf N), phosphorus (leaf P) and chlorophyll (Chl) and two physiological processes, viz., stomatal conductance (Gs net) and photosynthetic rate (A net), and RGR, of four dominant tree seedling species of a TDF (viz., Buchanania lanzan, Diospyros melanoxylon, Shorea robusta and Terminalia tomentosa) on four sites were analysed for species, site and season effects over a 2-year period. Step-wise multiple regression was performed to predict RGR from mean values of SMC, leaf traits and physiological processes. Principal component analysis (PCA) was performed to observe the extent of intra- vs. inter-specific variability in the leaf traits and physiological rates.Important findings All the traits and physiological rates were interrelated and showed significant positive relationship with RGR except for the correlation of LDMC with RGR which was not significant. Further, relationships of SMC with all leaf traits, physiological rates and RGR were significant, except for that between SMC and SLA for B. lanzan and D. melanoxylon. The slope of seedling trait:SMC relationship, a measure of phenotypic plasticity in response to soil moisture gradient, varied among species. Among the four species, T. tomentosa was the most plastic and S. robusta the least. In conclusion, leaf traits and physiological processes were strongly related to soil water availability on the one hand and seedling growth on the other. Gs net is the most important variable which accounted for the greatest amount of variability (62%) in RGR, emphasizing the role of stomatal conductance in shaping growth patterns across spatial and temporal gradients of soil water availability. Gs net and SMC together explained 64% variability in RGR, indicating that other traits/factors, not studied by us are also important in modulating the growth of tropical tree seedlings.     

Tree community structure and stem mortality along a water availability gradient in a Mexican tropical dry forest

We document spatial changes in species diversity, composition, community structure, and mortality of trees across a gradient of water availability in a tropical dry forest in western Mexico. This gradient occurs along the main stream of a small watershed of less than 1 km in length. Four 30 × 80 m plots were established systematically to include the driest (ridge top of the watershed) to the wettest sites (watershed bottom) within this watershed. All stems larger than 5 cm were identified, and measured for diameter and height. Dead stems larger than 5 cm were measured and classified as: a) found on live or dead trees, and b) standing (“snags”) or lying (“downlogs”) on the ground. The number of recorded species per plot declined from 73 to 44 species as water availability decreased. A decline in estimated total richness, and in Shannon-Wiener and Simpson diversity indices was also observed in the drier plots. Species composition strongly changed along the gradient, with the two ends of the gradient sharing only 11% of the species. Stem density and percentage of dead stems and trees increased in abundance and basal area from the wetter to the drier sites. Tree and stem size (basal area, height and stem diameter) showed the opposite trend. Nonetheless, total basal area of live trees was largest at the two end gradient locations and oscillated between 12.22 m² ha⁻¹ and 7.93 m² ha⁻¹. Proportion of snags increased towards the driest site (from 46 to 72%), while that of down logs decreased. Overall, our results suggest that small-scale gradients of water availability play a paramount role in the spatial organization of tree communities in seasonal tropical environments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Tree growth traits and social status affect the wood density of pioneer species in secondary subtropical forest

Wood density (WD) is not only an important parameter to estimate aboveground biomass but also an indicator of timber quality and plant adaptation strategies to stressful conditions (i.e., windthrow, pests, and pathogens). This study had three objectives: (1) to compare WD among seven subtropical tree species; (2) to determine how tree growth traits may influence possible differences in WD between the pioneer and shade‐tolerant species; and (3) to examine whether or not WD differs by tree social status (dominant vs. suppressed trees) within species. To do this, 70 trees were destructively harvested. From each tree, disks at different stem heights were obtained and subjected to a method of stem analysis to measure whole tree level WD. The results showed that WD differed significantly among the seven species (p < .001). Their average WD was 0.537 g/cm³, ranging from 0.409 g/cm³ for Choerospondias axillaris to 0.691 g/cm³ for Cyclobalanopsis glauca. The average WD of the four pioneer species (0.497 ± 0.13 g/cm³) was significantly lower (p < .01) than that of the three shade‐tolerant species (0.589 ± 0.12 g/cm³). The WD of the pioneers had a significant positive correlation with their stem diameter at breast height (DBH), tree height (H), and tree age, but WD had a significant negative correlation with relative growth rate (RGR). In contrast, the WD of the shade‐tolerant tree species had no significant relationships with DBH, H, tree age, or RGR. The dominant trees of the pioneer species had a higher WD than the suppressed trees, whereas the shade‐tolerant species had a lower WD for dominant trees than the suppressed trees. However, the differences in WD between dominant and suppressed trees were not significant. Taken together, the results suggest that classifying species into pioneer and shade‐tolerant groups to examine the effects of tree growth traits and social status could improve our understanding of intra‐ and interspecific variation in WD among subtropical tree species.     

Phosphorus cycling in a Mexican tropical dry forest ecosystem

The study was conducted in five contiguous small watersheds (12–28 ha) gauged for long-term ecosystem research. Five 80 × 30 m plots were used for the study. We quantified inputs from the atmosphere, dissolved and particulate-bound losses, throughfall and litterfall fluxes, standing crop litter and soil available P pools. Mean P input and output for a six-year period was 0.16 and 0.06 kgha^–1yr^–1, respectively. Phosphorus concentration increased as rainfall moved through the canopy. Annual P returns in litterfall (3.88 kg/ha) represented more than 90% of the total aboveground nutrient return to the forest floor. Phosphorus concentration in standing litter (0.08%) was lower than that in litterfall (0.11%). Phosphorus content in the litterfall was higher at Chamela than at other tropical dry forests. Mean residence time on the forest floor was 1.2 yr for P and 1.3 yr for organic matter. Together these results suggest that the forest at Chamela may not be limited by P availability and suggest a balance between P immobilization and uptake. Comparison of P losses in stream water with input rates from the atmosphere for the six-year period showed that inputs were higher than outputs. Balances calculated for a wet and a dry year indicated a small P accumulation in both years.     

Plant functional traits under cattle grazing and fallow age scenarios in a tropical dry forest of Northwestern Mexico

Evaluating plant functional traits helps to understand how plants respond to changing environmental conditions and resource availability associated with disturbance events. Livestock production is one of the primary drivers of tropical forest loss and degradation. Livestock alter environmental conditions within the forest by grazing, trampling and nutrient inputs, which in turn can influence species composition and functional traits of species. Understanding how livestock influence functional traits along a successional gradient is poorly understood. Here, we studied the effect of cattle grazing and fallow age on plant functional traits and soil nutrients in secondary and old-growth tropical dry forests. We analyzed plant functional traits of the most important species in successional and old-growth forest communities in both cattle present and cattle excluded plots. Our results showed the effects of cattle grazing and fallow age on plant functional traits, with fallow age explaining more variation than cattle grazing. In early succession, functional traits were associated with water conservation (thicker leaves, lower specific leaf area), and in later successional they were linked with sunlight conservation (larger height, higher specific leaf area). The presence of large fruits and seeds in advanced successional sites suggests high resource availability, which may help plants to successfully reproduce. Moreover, under cattle grazing some functional traits are associated with herbivory defense (high foliar dry weight and thick leaves). Even though N and C increased as succession advanced, the sites with cattle grazing had higher NH4 and NO3 concentrations as a result of fecal deposition. Plant functional traits responded to fallow age than to cattle grazing. Our study showed that cattle exclusion, as a management and biodiversity conservation strategy, contributes positively to soil nutrition. Thus, fallow age and cattle exclusion facilitate soil recovery and allows establishing species with suitable functional attributes for overcoming environmental filters in abandoned cattle fields.     

Seedling recruitment patterns in a tropical dry forest in Ghana

Abstract. Studies of seedling demography and dynamics were carried out in a dry forest at Pinkwae, Ghana (rainfall 1100 mm). Seedlings of tree and liana species in permanent transects were tagged, identified, measured and recensused at bimonthly intervals. A total of 1931 seedlings in 53 species were followed over 14 censuses during a 2-yr period. Germination and mortality were seasonal in their distribution; mortality was highest in dry periods. Seedling density varied seasonally, with the highest densities in comparatively wet periods. Diversity of seedlings was low, corresponding to that of the adult assemblage. Survivorship of tagged seedlings was recorded during the study period. The cohort tagged at the first census, comprising 1033 individuals of mixed ages and species, lost 61.9% during a 24-month period. Many seedlings were found to be rather large at the initial post-germination stage, with strong root development. Vegetative reproduction by root suckers or epicormic shoots was quite common at the site, although the majority of regeneration was by seeds. The current seedling assemblage appears to be representative, in terms of composition and diversity, of the adult species assemblage at Pinkwae.     

Hydraulic and photosynthetic co-ordination in seasonally dry tropical forest trees

In the present study the linkage between hydraulic, photosynthetic and phenological properties of tropical dry forest trees were investigated. Seasonal patterns of stem‐specific conductivity (K_SP) described from 12 species, including deciduous, brevi‐deciduous and evergreen species, indicated that only evergreen species were consistent in their response to a dry‐to‐wet season transition. In contrast, K_SP in deciduous and brevi‐deciduous species encompassed a range of responses, from an insignificant increase in K_SP following rains in some species, to a nine‐fold increase in others. Amongst deciduous species, the minimum K_SP during the dry season ranged from 6 to 56% of wet season K_SP, indicating in the latter case that a significant portion of the xylem remained functional during the dry season. In all species and all seasons, leaf‐specific stem conductivity (K_L) was strongly related to the photosynthetic capacity of the supported foliage, although leaf photosynthesis became saturated in species with high K_L. The strength of this correlation was surprising given that much of the whole‐plant resistance appears to be in the leaves. Hydraulic capacity, defined as the product of K_L and the soil–leaf water potential difference, was strongly correlated with the photosynthetic rate of foliage in the dry season, but only weakly correlated in the wet season.     

Tree species diversity driven by environmental and anthropogenic factors in tropical dry forest fragments of central Veracruz, Mexico

We examined vegetation structure and woody species diversity in relation to 14 environmental and anthropogenic factors in ten tropical dry forest (TDF) fragments in central Veracruz, Mexico. The basal area of the canopy (30.2 ± 2.11 m²/ha) and understory (1.96 ± 0.12 m²/ha) trees was similar, but density (1,014 ± 104 and 2,532 ± 227 individuals/ha, respectively) differed among sites. We recorded 98 canopy, 77 understory, and 60 seedling species. Richness was 24–45 species per site, Fisher’s alpha and Shannon’s indices increased with site altitude. Chao Jaccard indices revealed high species turnover, and a consistently higher similarity within the sites at the lowest and within the highest elevation sites. Ordination identified altitude, aspect, slope, water proximity, cattle and trails as significant explanatory variables of species patterns, and showed that sites at lower elevations were clearly separated from the other sites. Environmental heterogeneity alone did not control species diversity distribution, but species were affected by environmental filters at different stages in their life cycle, e.g., water proximity was significant for saplings and seedlings but not for adults. Anthropogenic disturbances act synergistically, e.g., trails played a key role in determining structure and tree diversity patterns. An important finding is that human disturbance diminishes species diversity in this TDF, but sites at lower elevations were more disturbed and less diverse, therefore we need to study how environmental factors would act if there were no anthropogenic disturbance.     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Root biomass of a dry deciduous tropical forest in Mexico

The deciduous tropical dry forest at Chamela (Jalisco, Mexico) occurs in a seasonal climate with eight rainless (November through June) and four wet months (700 mm annual precipitation). The forest reaches a mean height of 10 m. Tree density in the research area was 4700 trees per ha with a basal area at breast height of 23 m² per ha. The above-and below-ground biomass of trees, shrubs, and lianas was 73.6 Mg ha^–1 and 31 Mg ha^–1, respectively. A root:shoot biomass ratio of 0.42 was calculated. Nearly two thirds of all roots occur in the 0–20 cm soil layer and 29% of all roots have a diameter of less than 5 mm.     

Leaf attributes and tree growth in a tropical dry forest

Questions: How are leaf attributes and relative growth rate (RGR) of the dominant tree species of tropical deciduous forest (TDF) affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional attributes with each other? Can leaf attributes singly or in combination predict the growth rate of tree species of TDF? Location: Sonebhadra district of Uttar Pradesh, India. Methods: Eight leaf attributes, specific leaf area (SLA); leaf carbon concentration (LCC); leaf nitrogen concentration (LNC); leaf phosphorus concentration (LPC); chlorophyll concentration (Chl), mass‐based stomatal conductance (Gs_mass); mass based photosynthetic rate (A_mass); intrinsic water use efficiency (WUEi); and relative growth rate (RGR), of six dominant tree species of a dry tropical forest on four sites were analysed for species, site and season effects over a 2‐year period. Step‐wise multiple regression was performed for predicting RGR from mean values of SMC and leaf attributes. Path analysis was used to determine which leaf attributes influence RGR directly and which indirectly. Results: Species differed significantly in terms of all leaf attributes and RGR. The response of species varied across sites and seasons. The attributes were positively interrelated, except for WUEi, which was negatively related to all other attributes. The positive correlation was strongest between Gs_mass and A_mass and the negative correlation was strongest between Gs_mass and WUEi. Differences in RGR due to site were not significant when soil moisture was controlled, but differences due to season remained significant. The attributes showed plasticity across moisture gradients, which differed among attributes and species. Gs_mass was the most plastic attribute. Among the six species, Terminalia tomentosa exhibited the greatest plasticity in six functional attributes. In the step‐wise multiple regression, A_mass, SLA and Chl among leaf attributes and SMC among environmental factors influenced the RGR of tree species. Path analysis indicated the importance of SLA, LNC, Chl and A_mass in determining RGR. Conclusion: A _mass, SMC, SLA and Chl in combination can be used to predict RGR but could explain only three‐quarters of the variability in RGR, indicating that other traits/factors, not studied here, are also important in modulating growth of tropical trees. RGR of tree species in the dry tropical environment is determined by soil moisture, whereas the response of mature trees of different species is modulated by alterations in key functional attributes such as SLA, LNC and Chl.     

Limnological observations of an intermittent tropical dry forest stream

This study examines aspects of the seasonal limnology of an intermittent stream in the dry forest region of northwest Costa Rica. It focuses on annual water level fluctuations and both seasonal and among pool variation in dissolved oxygen concentration and water temperature. Dry season pools differed in morphometry and the rate of decline in water levels subsequent to the seasonal floods. Rate of water level decline was related both to pool depth and to the exposure of the pool to the sun. Oxygen concentration was generally low in the dry season, but increased during the rainy season in association with rain events and seasonal flooding. A repeated-measures analysis of variance indicated that seasonal effects accounted for 40% of the variance in oxygen concentration for residual pools. Differences among pools were also significant. However, there was strong evidence for a significant interaction between seasonal and spatial influences on oxygen values in the system. Variation in water temperature was small, but differences among pools, sampling dates, and their interaction were all significant. Periphyton production increased significantly between the late wet season sample in November and the dry season sample in February. Incident light intensity explained 76% of the variation among pools in net periphyton production in the dry season.     

Dispersal modes affect tropical forest assembly across trophic levels

We examined assemblages of trees and two major groups of vertebrate seed dispersers, birds and primates, in Ugandan protected areas to evaluate the roles of dispersal limitation and species sorting in community assembly. We conducted partial Mantel tests to investigate relationships between community similarity, environmental distance and geographic distance. Results showed that environmental factors, specifically temperature and rainfall, significantly and more strongly structured tree assemblages than geographic distance. Analysis of tree dispersal modes revealed wind‐dispersed tree guilds were significantly dispersal limited but trees dispersed by animals were not. For assemblages of vertebrate seed dispersers, dispersal limitation significantly and more strongly structured assemblages of primates than species sorting whereas environmental factors significantly and more strongly structured assemblages of birds than dispersal limitation. We therefore examined whether trees dispersed by primates were more dispersal limited than trees dispersed by birds. We found consistent trends that primate fruit trees were more dispersal limited than bird fruit trees using three definitions of dispersal syndromes based on fruit color. Our results suggest that the dispersal abilities of primary consumers may affect the distribution of primary producers at large spatial scales.     

Environmental control of flowering periodicity in Costa Rican and Mexican tropical dry forests

Aim We analyse the proximate causes of the large variation in flowering periodicity among four tropical dry forests (TDF) and ask whether climatic periodicity or biotic interactions are the ultimate causes of flowering periodicity. Location The four TDFs in Guanacaste (Costa Rica), Yucatan, Jalisco and Sonora (Mexico) are characterized by a 5–7 month long dry season and are located along a gradient of increasing latitude (10–30°N). Methods To dissect the differences in flowering periodicity observed at the community level, individual tree species were assigned to ‘flowering types’, i.e. groups of species with characteristic flowering periods determined by similar combinations of environmental flowering cues and vegetative phenology. Results Large variation in the fraction of species and flowering types blooming during the dry and wet season, respectively, indicates large differences in the severity of seasonal drought among the four forests. In the dry upland forests of Jalisco, flowering of leafless trees remains suppressed during severe seasonal drought and is triggered by the first rains of the wet season. In the other forests, leaf shedding, exceptional rainfall or increasing daylength cause flowering of many deciduous species at various times during the dry season, well before the summer rains. The fraction of deciduous species leafing out during the summer rains and flowering when leafless during the dry season is largest in the Sonoran TDF. Main conclusions In many wide‐ranging species the phenotypic plasticity of flowering periodicity is large. The distinct temporal separation of spring flowering on leafless shoots and subsequent summer flushing represents a unique adaptation of tree development to climates with a relatively short rainy season and a long dry season. Seasonal variation in rainfall and soil water availability apparently constitutes not only the proximate, but also the ultimate cause of flowering periodicity, which is unlikely to have evolved in response to biotic adaptive pressures.