The role of canopy gaps in the regeneration of coastal dune forest

In regenerating coastal dune forest, the canopy consists almost exclusively of a single species, Acacia karroo. When these trees die, they create large canopy gaps. If this promotes the persistence of pioneer species to the detriment of other forest species, then the end goal of a restored coastal dune forest may be unobtainable. We wished to ascertain whether tree species composition and richness differed significantly between canopy gaps and intact canopy, and across a gradient of gap sizes. In three known‐age regenerating coastal dune forest sites, we measured 146 gaps, the species responsible for gap creation, the species most likely to reach the canopy and the composition of adults, seedlings and saplings. We paired each gap with an adjacent plot of the same area that was entirely under intact canopy and sampled in the same way. Most species (15 of 23) had higher abundance in canopy gaps. The probability of self‐replacement was low for A. karroo even in the largest gaps. Despite this predominance of shade‐intolerant species, regenerating dune forest appears to be in the first phase of succession with ‘forest pioneers’ replacing the dominant canopy species. The nature of these species should lead to successful regeneration of dune forest.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Tree species regeneration in a mid-elevation,temperate rain forest in Isla de Chiloé, Chile

The regeneration of canopy and subeanopy species in a mid-elevation, primary rain forest in the Coastal Range of Isla de Chiloé (42°30S), in the cold-temperate region of Chile, was studied by comparing seedling and sapling abundances under the forest canopy, and within 36 tree-fall gaps. The forest was dominated byAmomyrtus luma andLaurelia philippiana (33 and 32% of the main canopy individuals), and two subcanopy species (Myrceugenia ovata, andMyrceugenia planipes) were also important. Uncommon species in the canopy wereDrimys winteri, Amomyrtus meli, andRaphithamnus spinosus. Tree-fall gaps were created generally by the fall of several trees, and the main canopy species were the principal gap-makers. Gap sizes varied between 28 and 972 m², with a mean of 197 m². Seedling and sapling abundances indicate that the dominant species are capable of regenerating below the canopy, but they also germinate and show enhanced growth within small light gaps. For one of the common subcanopy species (M. planipes) and the two infrequent canopy species (D. winteri, andA. meli) regeneration seems to depend entirely on tree-fall gaps. Thus, in this forest, light gaps allow the persistence of infrequent canopy species, but seem less important for the regeneration and maintenance of dominant canopy species.     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

Dynamics of forests with Araucariaceae in the western Pacific

Abstract. Several species of Araucaria and Agathis (Araucariaceae) occur as canopy emergents in rain forests of the western pacific region, often representing major components of total stand biomass. New data from permanent forest plots (and other published work) for three species (Araucaria hunsteinii from New Guinea, A. laubenfelsii from New Caledonia, and Agathis australis from New Zealand) are used to test the validity of the temporal stand replacement model proposed by Ogden (1985) and Ogden & Stewart (1995) to explain the structural and compositional properties of New Zealand rain forests containing the conifer Agathis australis. Here we propose the model as a general one which explains the stand dynamics of rain forests with Araucariaceae across a range of sites and species in the western Pacific. Forest stands representing putative stages in the model were examined for changes through time in species recruitment, growth and survivorship, and stand richness, density and basal area. Support for the model was found on the basis of: 1. Evidence for a phase of massive conifer recruitment following landscape-scale disturbances (e.g. by fire at the Huapai site, New Zealand for Agathis australis); 2. Increasing species richness of angiosperm trees in the pole stage of forest stand development (i.e. as the initial cohort of conifers reach tree size; >10 cm DBH); 3. A high turnover rate for angiosperms (<100 yr), and low turnover for conifers (≥ 100 yr) in the pole stage, but similar turnover rates for both components (50–100 yr) as forests enter the mature to senescent phase for the initial conifer cohort; 4. Very low rates of recruitment for conifers within mature stands, and projected forest compositions which show increasing dominance by angiosperm tree species; 5. A low probability of conifer recruitment in large canopy gaps created by conifer tree falls during the initial cohort senescent phase, which could produce a second generation low density stand in the absence of landscape scale disturbance; 6. Evidence that each of the three species examined required open canopy conditions (canopy openness > 10 %) for successful recruitment. The evidence presented here supports the temporal stand replacement model, but more long-term supporting data are needed, especially for the phase immediately following landscape level disturbance.     

Small gap dynamics in the southern boreal forest of eastern Canada: Do canopy gaps influence stand development?

Question: To what extent do small‐scale disturbances in the forest canopy, created by natural disturbance agents, affect stand development? Doubts exist as to whether small canopy openings have any real effect on the understory tree recruitment, especially in boreal forests. Location: Conifer and mixed stands in the Gaspesian region in eastern Québec. The main natural disturbance agents are recurring outbreaks of Choristoneura fumiferana (eastern spruce budworm) and winds. Methods: Linear transects in 27 sites were used to describe the gap (< 0.1 ha) regime parameters, including gap fraction, gap size and change in disturbance severity through time. Three stand types were distinguished, based on a gradient of abundance of tree host species for the eastern spruce budworm. The impact of gaps was evaluated on the basis of changes in the number, the period of recruitment, and the composition of tree saplings present within gap areas. Changes were measured along the gap size gradient, and according to the pattern of recent budworm epidemics. Results: The gap fraction is highly variable (18%‐64%) and is on average relatively high (42%). Gap sizes have a positively skewed distribution. In most cases the growth rate among gap filling saplings increased sufficiently to date disturbance events. The composition and the structure of understory trees were affected by gap formation. The number of shade‐intolerant tree species did increase during or following periods of particularly severe canopy disturbances. However, the establishment or survival of shade intolerant species was not restricted to larger gaps or more intensely disturbed periods. Conclusions: In sub‐boreal forests of Eastern Canada, small scale disturbances in the tree canopy influence stand regeneration dynamics, but not to the extent that parameters such as sapling composition and recruitment patterns depend on gap regime characteristics.     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Factors influencing sapling composition in canopy gaps of a temperate deciduous forest

To detect the factors that affect sapling species composition in gaps, we investigated 55 gaps in an old-growth temperate deciduous forest in Ogawa Forest Reserve, central Japan. Gap size, gap age, gap maker species, topographic location, adult tree composition around gaps, and saplings of tree species growing in the gaps were censused. For gaps 5 m², mean gap size was 70 m² and the maximum was 330 m². Estimated ages of gaps had a tendency to be concentrated in particular periods relating to strong wind records in the past. The sapling composition in gaps was highly and significantly correlated to that under closed canopy, indicating the importance of advance regeneration in this forest. However, some species showed significant occurrence biases in gaps or under closed canopy, suggesting differences in shade tolerance. The result of MANOVA showed that gap size and topography were important factors in determining the sapling composition in gaps. Species of gap makers affected the sapling composition indirectly by influencing gap size. The existence of parent trees around gaps had effects on sapling densities of several species. Gap age did not have clear influences on sapling composition. Variations in gap size and topography were considered as important factors that contribute to maintenance of species diversity in this forest.     

Shoot growth and tree architecture of saplings of the major canopy dominants in a warm-temperate rainforest

Sapling density, shoot growth, and sapling architecture were studied in five major canopy dominants both under closed canopy and gaps in a warm-temperate rainforest. The five species showed wide variations in distribution, shoot growth, and sapling architecture. Distylium racemosum and Quercus acuta had significantly higher sapling densities under closed canopy than in gaps. Castanopsis sieboldii and Machilus thunbergii had significantly higher sapling densities in gaps than under closed canopy. Quercus salicina showed no significant difference in sapling density between the two habitats. Under closed canopy, C. sieboldii and M. thunbergii had wider crowns than the other species. Distylium racemosum had the greatest number of terminal shoots among the species. Quercus acuta had a branchless small crown. Quercus salicina showed intermediate values in crown width, depth and the number of terminal shoots among the species. Distylium racemosum showed the greatest height-growth rate among the species under the closed canopy, but was the slowest in gaps. Castanopsis sieboldii and M. thunbergii showed the greatest height-growth rates among the species in gaps. Quercus salicina showed the slowest height-growth rates both under closed canopy and gaps. All of the five species showed low mortality under closed canopy. For the major canopy dominants: (i) sapling architecture may not be an important factor in determining mortality but it may be important for height-growth rate; and (ii) sprouting helps saplings to survive until gap formation and facilitates rapid growth in the gaps.     

Extant and potential vegetation of an old-growth maritime Ilex opaca forest

The extant and potential (seed bank) vegetation of a rare maritime holly forest on Fire Island, New York was described to assess whether treefall gaps act as a mechanism for the persistence of the species composition of this plant community over time. The Sunken Forest overstory is dominated by Ilex opaca, Amelanchier canadensis and Sassafras albidum. A survey of canopy gaps indicated canopy openings compose 11.3% of the land within the Sunken Forest (16 ha). The composition and density of the seed bank were described using the emergence method. Germination from soil samples placed in the greenhouse was monitored over 2 years. Sixteen species germinated with an average propagule density of 215±41 germinants per square metre. An early successional species (Rhus copallinum) dominated the seed bank, but the late-successional, shade-tolerant I. opaca was also present. Though only one species in the seed bank did not appear in the current vegetation, species abundance differed between vegetation strata. The mean cover and density of the ground-layer flora were higher beneath treefall gaps than closed canopy. Sapling density did not differ between the two canopy conditions, but the dominant species differed with A. canadensis occupying several closed canopy plots and P. serotina saplings appearing more often in gap plots. Most of the dominant canopy species are present in the seed bank and ground layer but are not present in the shrub and sapling layer, with the exception of A. canadensis. Current (2002) sapling density is lower than three decades ago for all species except P. serotina, which is now the dominant woody species in the Sunken Forest understory. The results of this study indicate that if the cause of the sapling reduction is lessened or removed, the characteristic species of the overstory of this unusual plant community may rebound and redevelop a sapling and shrub layer akin to that present before the increase in Odocoileus virginianus on the island.     

Stand dynamics during a 12 year period in an old-growth,cool temperate forest in northern Japan

In 1979 and 1991, trees over 2.0 m high were measured and mapped together with their crown projections to clarify stand dynamics and shifts in canopy dominants during this period, in a permanent plot of 0.525 ha in an old-growth, cool temperate mixed forest of Mt Moiwa, Central Hokkaido, northern Japan. During this period, an abundant recruitment of trees was observed after some canopy trees were felled by a typhoon in 1981 leaving gaps in the canopy. Vigorous recruitment was observed forTilia japonica, Acer mono andPrunus ssiori. These species had different regeneration sites in relation to canopy state. NeitherUlmus japonica norKalopanax pictus had any recruits during the 12 year period even in gaps. The equilibrium composition of tree species projected from transition probability analysis also implied the above shift of dominant species during the 12 year period in the plot and suggested that the present forest is not in an equilibrium state.     

Seed shadow, seedling recruitment, and spatial distribution of Buchenavia capitata (Combretaceae) in a fragment of the Brazilian Atlantic forest.

Here we describe the seed shadow, seedling recruitment, ontogenetic structure and spatial distribution of Buchenavia capitata (an emergent canopy tree) in a 380-ha fragment of the Atlantic forest in northeast Brazil. In particular, we examine seed distribution around 10 parental trees and both seedling recruitment and mortality, during an 18 month period beneath and around parental trees. Moreover, we describe: (1) B. capitata occurrence within treefall gaps; (2) population structure in terms of ontogenetic stages for the whole site; and (3) spatial distribution of adults within an area of 51 hectares. 99% of seeds were found beneath parent crowns (n = 4,236) and seed density reached 14.6 +/- 29.9 seeds/m2 (0-140 seeds/m2). 49% of all seeds germinated but seedling mortality reached 100% after an 18 month period. In addition, saplings of B. capitata were not found in forest understory and within 30 treefall gaps (94-2,350 m2). The adults showed an average DBH of 69.3 +/- 22.1 cm, were 19.2 +/- 2.9 m tall and presented a clumped spatial distribution. B. capitata matched some of the features presented by shade intolerant trees or large-gap specialists, and we hypothesize that low rates or even lack of long distance seed dispersal events may be reducing the probability of B. capitata seeds reaching suitable habitats for successful seedling recruitment and growth. Because of that (1) seedlings face high levels of early mortality; (2) there is no sapling recruitment at the study site; and (3) local population faces senility and it is threatened by local extinction.     

Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest in Aya district,southwestern Japan

The population structure and regeneration of canopy species were studied in a 4 ha plot in an old-growth evergreen broad-leaved forest in the Aya district of southwestern Japan. The 200 m × 200 m plot contained 50 tree species, including 22 canopy species, 3,904 trees (dbh5 cm) and a total basal area of 48.3 m²/ha. Forty one gaps occurred within the plot, and both the average gap size (67.3 m²) and the total area of gap to plot area (6.9%) were small. Species found in the canopy in the plot were divided into three groups (A, B, C) based on size and spatial distribution patterns, and density in each tree size. Group A (typical species: Distylium racemosum, Persea japonica) showed a high density, nearly random distribution and an inverse J-shaped size distribution. Species in group B (Quercus salicina, Quercus acuta, Quercus gilva) were distributed contagiously with conspicuous concentration of small trees (<5 cm dbh) around gaps. However, the species in this group included few trees likely to reach the canopy in the near future. Group C included fast-growing pioneer and shade intolerant species (e.g. Cornus controversa, Carpinus tschonoskii, Fagara ailanthoides), which formed large clumps. Most gaps were not characterized by successful regeneration of group B and C but did appear to accelerate the growth of group A. Group B species appear to require long-lived or large gaps while group C species require large, catastrophic disturbances, such as landslides, for regeneration.     

Community dynamics of evergreen broadleaf forests in southwestern Japan

The process and rate of revegetation in gaps in an evergreen oak forest were studied by comparing the species composition, tree density, frequency distribution of tree height, and relation between diameter at breast height and tree height among different aged stands. For estimating stand ages, the ages of gap indicators, such as,Symplocos prunifolia andAcer rufinerve, were very useful. It took about 70 years for gaps to be filled by large fully-grown trees. Since the mean residence time of the forest canopy was 180 years, the trees that attain the forest canopy were expected to be canopy trees for 110 years on the average. Tree densities of all broadleaved evergreens exceptS. prunifolia, were independent of stand age. On the other hand, densities of gap indicators,S. prunifolia andA. rufinerve, decreased as stand age increased. Other deciduous broadleaf and coniferous species were scarce as a whole. According to the frequency distributions of height of live and dead trees in different aged stands, it was suggested that shorter trees were more susceptible to death than taller trees. The self-thinning in revegetation process in gaps approximately followed the 3/2 power law, though the power was larger (−1.32) than expected from the law.     

Recruitment patterns and northward tree migration through gap dynamics in an old-growth white pine forest in northern Ontario

Decreases in abundances and declines in growth of eastern white pine over the past century due mainly to human activities have resulted in few large intact old-growth white pine forests in Ontario. These stands may be vulnerable to replacement by deciduous species from temperate forests further south, where recruitment in canopy gap disturbances can greatly define the regeneration process. We investigated recruitment dynamics in canopy gaps of an old-growth white pine forest of Temagami, northern Ontario, Canada, the northern limit of the temperate?Cboreal ecotone. White pine, red pine, black spruce and eastern white cedar represented 85?% of the mature canopy abundance, where trees and saplings established equally in gaps and the closed canopy. Balsam fir and paper birch were more abundant in gaps, showing increases of abundance and basal area with increases in gap size representing canopy self-replacement (balsam fir) and autogenic succession (paper birch). Red maple, at its northernmost range limit, was the only species to show linear increases of abundance and basal area with increases in gap size and gap age. This result, along with adult red maples present in gaps but absent from the closed canopy, identifies the establishment of a northward migrating species in gaps as hypothesized for pine forests at the northern limit of this broad ecotone. We discuss how migration pressures, coupled with pine recruitment limitation through reduced fire frequency by regional fire suppression and predicted future increased warming of 2?C4?°C over the next century, threatens replacement of old-growth white pine forests at this latitude with northward migrating tree species found further south.     

Environmental Correlates of Tree and Seedling–Sapling Distributions in a Mexican Tropical Dry Forest

Bray and Curtis ordination was used to explore which environmental variables explained importance values and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra de Manantlán, Jalisco, Mexico. The diameters of trees ≥2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis ordination. The matrices were based on density, frequency, and dominance of adult trees as well as seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope, canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1 and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling presence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization along different soil microsites could promote species coexistence in this forest, while heterogeneity in light conditions may instead determine differences in growth and, thus, importance value of trees. We hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely more importance at early stages in tree life histories than in later life history.     

Patch structure and ramet demography of the clonal tree, Asimina triloba, under gap and closed-canopy

Clonal understory trees develop into patches of interconnected and genetically identical ramets that have the potential to persist for decades or centuries. These patches develop beneath forest canopies that are structurally heterogeneous in space and time. Canopy heterogeneity, in turn, is responsible for the highly variable understory light environment that is typically associated with deciduous forests. We investigated what aspects of patch structure (density, size structure, and reproductive frequency of ramets) of the clonal understory tree, Asimina triloba, were correlated with forest canopy conditions. Specifically, we compared A. triloba patches located beneath closed canopies and canopy gaps. We also conducted a three-year demographic study of individual ramets within patches distributed across a light gradient. The closed canopy-gap comparison demonstrated that the patches of A. triloba had a higher frequency of large and flowering ramets in gaps compared to closed-canopy stands, but total ramet density was lower in gaps than in closed canopy stands. In the demographic study, individual ramet growth was positively correlated with light availability, although the pattern was not consistent for all years. Neither ramet recruitment nor mortality was correlated with light conditions. Our results indicate that the structure of A. triloba patches was influenced by canopy condition, but does not necessarily depend on the responses of ramets to current light conditions. The lack of differences in ramet recruitment and mortality under varying canopy conditions is likely to be a primary reason for the long-term expansion and persistence of the patches. The primary benefit of a positive growth response to increasing light is the transition of relatively small ramets into flowering ramets within a short period of time.