Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

The interacting effects of herbivore exclosures and seed addition in a wet meadow

Seed limitation may prevent successful restoration of native plant communities. Seed addition is a common restoration practice but the role of small mammals in affecting seedling recruitment is not well understood. The purpose of this investigation was to test the relative effect of seed introduction in combination with small mammal and bird exclosures in an Ohio wet meadow. We ask whether the ambient population of Microtus pennsylvanicus (1) alters species composition (e.g. forb/grass/sedge, invasive, non-native); (2) influences plant diversity; and (3) reduces the effect of increasing local plant richness through seed introductions. We established a 2×2 factorial design including a seed addition treatment (0 and 20 seed species added) and an exclosure treatment (open and fenced to exclude all mammalian and bird herbivores and granivores). Seeds from twenty native species were selected to represent a broad range of plant life forms typically found in temperate eastern North American wet meadow communities. All species were obligate or facultative wetland species with forbs, grasses and sedges represented. We found that forb species increased inside exclosures, especially in the seed addition treatment. We also found that relative biomass of invasive species was reduced in exclosures and with seed addition. Species richness increased with seed addition; however, exclosures significantly increased species richness and diversity, particularly of those species that were experimentally introduced by seed. Our results support the seed limitation hypotheses. It is also evident that seed and seedling predation are important factors that can control wet meadow community composition and diversity.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Succession and Micro-elevation Effects on Seedling Establishment of Calophyllum brasiliense Camb. (Clusiaceae) in an Amazonian River Meander Forest1

I investigated the effects of successional stage and micro‐elevation on seedling establishment of Calophyllum brasiliense (Clusiaceae), a common canopy tree of seasonally flooded lowland forest along the Manú River meander zone in southeastern Peru. To compare seedling establishment between microhabitat types, I planted C. brasiliense seeds in a fully crossed experimental design of three successional stages (early, mid, and mature) and two micro‐elevations (levees and backwaters). Seedling establishment success in this study was affected by both successional stage and micro‐elevation, but micro‐elevation was most important in mid‐successional habitats. In general, seedlings in early succession experienced better conditions than in mature forest; light levels were higher, herbivory lower, and seedling growth higher. In mid‐successional forest, micro‐elevation determined habitat quality; backwaters had higher light levels, lower herbivory, and higher seedling growth and survival than levees. Mid‐successional backwaters were similar in quality to early successional forest for seedling establishment, while levees in that same successional stage were the poorest microhabitats for establishment. Although mid‐successional backwaters are similar to early succession for seedling establishment, in the long run, seedlings that establish in mid‐succession have a lower chance of reaching reproductive size before their habitat ages to mature forest than members of their cohort that established in early succession. I hypothesize that successful recruitment for C. brasiliense in the Manú River meander system requires dispersal to early successional habitat.     

The potential of small exclosures in assisting regeneration of coffee shade trees in South‐Western Ethiopian coffee forests

Ethiopian Afromontane moist forests where coffee grows as understorey shrub are traditionally managed by the local communities for coffee production through thinning of the shade tree canopy and slashing of competing undergrowth. This management practice has a negative impact on the coffee shrubs, because the removal of shade tree saplings and seedlings reduces the succession potential of the shade tree canopy, which threatens the very existence of the shade coffee production system. We assessed the functionality of small exclosures to initiate coffee shade tree canopy restoration through natural regeneration. Our results show that small exclosures have a strong restoration potential for the coffee shade trees preferred by farmers (Albizia schimperiana, A. gummifera and Millettia ferruginea), as evidenced from their seedling abundance, survival and growth. The regeneration of late‐successional tree species of the moist Afromontane forest was not successful in the small exclosures, most probably due to the low abundance or absence of adult trees as seed sources for regeneration. Therefore, temporary establishment of small exclosures in degraded coffee forest fragments where shade trees are getting old or dying is recommended for sustainable shade coffee production.     

An experimental test of density- and distant-dependent recruitment of mahogany (Swietenia macrophylla) in southeastern Amazonia

According to the Janzen–Connell model, high mortality of seeds and seedlings in proximity to conspecific adults can help maintain species diversity in tropical forests. Using a natural population of big-leaf mahogany (Swietenia macrophylla King), we tested the model’s mechanism by examining seed predation and juvenile recruitment in the forest understory and in treefall gaps in the vicinity of both isolated and clumped adults. We used tethered seeds placed in three types of exclosure plots: (1) complete access to seeds, (2) semi-access (access by small-sized seed predators) and (3) no access (all mammals excluded). Exclosure treatments were applied within the understory (both near and far from adults) and in gaps at eight fruiting adults in the late dry season (2001) and scored ten months later. Significantly more seeds were removed in canopy gaps near clumped adults than at isolated adults; otherwise, none of the treatment factors significantly influenced seed predation. In contrast, understory juvenile recruitment was significantly enhanced by distance from adults and was twice as high at isolated than clumped adults, providing novel support for the Janzen–Connell mechanism. No-access exclosures protected significantly more seeds than semi- and full-access exclosures, implicating small mammals in seed losses. Across the eight trees, juvenile recruitment in the no-access exclosures decreased significantly with conspecific adult densities, implicating non-mammalian density-responsive factor(s) in mortality following germination; likely a known specialist invertebrate herbivore. When all treatments were combined, conspecific adult basal area and total DBH explained 72 and 90% of variation in overall juvenile recruitment, respectively. Collectively, these results indicate that Janzen–Connell effects can operate in S. macrophylla, especially during the seed-to-seedling transition, and will likely reduce recruitment in areas of high conspecific densities. They also suggest that further research into the causes of density-dependence in tropical trees should investigate mortality agents following germination.     

Seedling responses to decreased snow depend on canopy composition and small‐mammal herbivore presence

Winter is becoming warmer and shorter across the northern hemisphere, and reductions in snow depth can decrease tree seedling survival by exposing seedlings to harmful microclimates. Similarly, herbivory by small mammals can also limit the survival and distribution of woody plants, but it is unclear whether winter climate change will alter small‐mammal herbivory. Although small‐scale experiments show that snow removal can either increase or decrease both soil temperatures and herbivory, we currently lack snow‐removal experiments replicated across large spatial scales that are needed to understand the effect of reduced snow. To examine how winter herbivory and snow conditions influence seedling dynamics, we transplanted Acer saccharum and Tsuga canadensis seedlings across a 180 km latitudinal gradient in northern Wisconsin, where snow depth varied seven‐fold among sites. Seedlings were transplanted into one of two herbivory treatments (small‐mammal exclosure, small‐mammal access) and one of two late‐winter snow removal treatments (snow removed, snow unmanipulated). Snow removal increased soil freeze‐thaw frequency and cumulative growing degree‐days (GDD), but the magnitude of these effects depended on forest canopy composition. Acer saccharum survival decreased where snow was removed, but only at sites without conifers. Excluding small mammals increased A. saccharum survival at sites where the small‐mammal herbivore Myodes gapperi was present. Excluding small mammals also increased T. canadensis survival in plots with < 5 cm snow. Because variation in canopy composition and M. gapperi presence were important predictors of seedling survival across the snow‐depth gradient, these results reveal complexity in the ability to accurately predict patterns of winter seedling survival over large spatial scales. Global change scenarios that project future patterns of seedling recruitment may benefit from explicitly considering interactions between snow conditions and small‐mammal winter herbivory.     

Small rodents as significant dispersers of tree seeds in a Neotropical forest

Abstract. Through seed dispersal and predation, terrestrial mammals should be an important component of the mechanisms that determine patterns of tree recruitment in tropical forests. Despite their great abundance and ubiquity in Neotropical forests, small rodents as seed predators and dispersers remain largely forgotten. To investigate the fates of seeds in a hunted primary forest in Belize, we tagged seeds of Astrocaryum mexicanum (Palmae), Ampelocera hottlei (Ulmaceae), and Pouteria sapota (Sapotaceae) and placed them into open plots, exclosures accessible only to small mammals, and exclosures accessible to medium-sized and small mammals. The exclosure experiments and fates of the seeds show that the spiny pocket mouse, Heteromys desmarestianus (Heteromyidae), was the dominant handler of seeds of the first two species and also removed a significant proportion of the very large-seeded Pouteria. Most of the seeds were killed immediately upon removal, but many of the seeds (3–18 %) of the first two species were scatterhoarded (dispersed and buried in the soil) by Heteromys. Some of the scatterhoarded seeds (29%) remain buried and therefore protected from predation by other animals. Agoutis (Dasyprocta punctata), a caviomorph rodent, buried 13 % of the seeds of Pouteria, and Heteromys consumed and dispersed but did not bury Pouteria seeds. Results of this study support predictions by some researchers that small rodents are dominant terrestrial granivores in Neotropical forests. The role of small rodents as seed dispersers, however, has never been fully appreciated.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

Enemies mediate distance- and density-dependent mortality of tree seeds and seedlings: a meta-analysis of fungicide,insecticide and exclosure studies

Conspecific negative distance- and density-dependence is often assumed to be one of the most important mechanisms controlling forest community assembly and species diversity globally. Plant pathogens, and insect and mammalian herbivores, are the most common natural enemy types that have been implicated in this phenomenon, but their general effects at different plant life stages are still unclear. Here, we conduct a meta-analysis of studies that involved robust manipulative experiments, using fungicides, insecticides and exclosures, to assess the contributions of different natural enemy types to distance- and density-dependent effects at seed and seedling stages. We found that distance- and density-dependent mortality caused by natural enemies was most likely at the seedling stage and was greater at higher mean annual temperatures. Conspecific negative distance- and density-dependence at the seedling stage is significantly weakened when fungicides were applied. By contrast, negative conspecific distance- and density-dependence is not a general pattern at the seed stage. High seed mass reduced distance- and density-dependent mortality at the seed stage. Seed studies excluding only large mammals found significant negative conspecific distance-dependent mortality, but exclusion of all mammals resulted in a non-significant effect of conspecifics. Our study suggests that plant pathogens are a major cause of distance- and density-dependent mortality at the seedling stage, while the impacts of herbivores on seedlings have been understudied. At the seed stage, large and small mammals, respectively, weaken and enhance negative conspecific distance-dependent mortality. Future research should identify specific agents of mortality, investigate the interactions among different enemy types and assess how global change may affect natural enemies and thus influence the strength of conspecific distance- and density-dependence.     

Diplochory in western chokecherry: you can��t judge a fruit by its mesocarp

Western chokecherry (Prunus virginiana var. demissa, Rosaceae) is dispersed by frugivorous birds and carnivores, but it has large seeds that are potentially attractive to rodents that could act as seed predators and dispersers. Here, we quantify the benefits of primary dispersal by birds and secondary dispersal by scatter-hoarding rodents. In the fall, avian frugivores (mostly American robins, Turdus migratorius, and cedar waxwings, Bombycilla cedrorum) consumed 87% of the fruit crop and dispersed 67% of the fruit crop away from parent plants. Rodents removed 89% of seeds that simulated bird-dispersed seed rain from transects in riparian zones and 58% from transects in upland habitats. Rodents scatter-hoarded 91.6% of the seeds they removed, burying most in small caches (two to eight seeds) 8?C25?mm deep. About 39% of the seeds in spring caches produced seedlings. Inside rodent-proof exclosures, 52.1% of seeds buried to simulate rodent caches produced seedlings, 29.7% of which were still alive after 1?year. In contrast, only 3.8% of seeds placed on the soil surface, simulating dispersal by avian frugivores, produced seedlings. Seed dispersal by frugivorous birds likely contributes to colonization of unoccupied habitat through long-range dispersal and to escape from distance-dependent seed mortality near the parent plant. Despite seed losses, rodents offer short-range seed dispersal and bury seeds in more favorable sites for germination, improving seedling emergence and establishment. The combined mechanisms of seed dispersal significantly enhanced chokecherry seedling recruitment by providing more dispersal-related benefits than either frugivorous bird or scatter-hoarding rodents could provide alone.     

Effects of herbivory and patch size on tree seedling survivorship in a fog-dependent coastal rainforest in semiarid Chile

The landscape (matrix) surrounding habitat fragments critically affects the biodiversity of those fragments due to biotic interchange and physical effects. However, to date, there have been only a limited number of studies on plant–animal interactions in fragmented landscapes, particularly on how tree seedling herbivory is affected by fragmentation. We have examined this question in a fog-dependent mosaic of rainforest fragments located on coastal mountaintops of semiarid Chile (30°S), where the effects of the surrounding semiarid matrix and forest patch size (0.1–22 ha) on tree seedling survival were simultaneously addressed. The rainforest is strongly dominated by the endemic evergreen tree species Aextoxicon punctatum (Olivillo, approx. 80% of basal area). To assess the magnitudes and causes of Olivillo seedling mortality, we set up a field experiment where 512 tree seedlings of known age were transplanted into four forest fragments of different sizes in four 1.5 × 3-m plots per patch; one-half of each plot was fenced off with chicken wire to exclude small mammals. The plots were monitored for 22 months. Overall, 50% of the plants died during the experiment. The exclusion of small mammals from the plots increased seedling survival by 25%, with the effect being greater in smaller patches where matrix-dwelling herbivores are more abundant. This experiment highlights the important role of the surrounding matrix in affecting the persistence of trees in forest fragments. Because herbivores from the matrix cause greater tree seedling mortality in small patches, their effects must be taken into account in forest conservation–restoration plans.     

An experimental study of the impacts of understorey forest vegetation and herbivory by red deer and rodents on seedling establishment and species composition in Waitutu Forest, New Zealand

Introduced mammalian herbivores are changing the structure and composition of New Zealand’s forest ecosystems and may modify forest succession after natural disturbances. We studied how introduced ungulates (red deer and feral pigs) and rodents (rats and house mice) affected the rate of recovery (i.e. the engineering resilience) of the forest understorey following artificial disturbance. We imposed disturbances by clearing understorey vegetation dominated by Blechnum ferns in forests on relatively fertile alluvium and elevated infertile marine terraces, and recorded recovery of vegetation (seedling establishment, species composition, cover and volume) in herbivore exclosures and controls. Seedlings quickly established on cleared plots: after 2 years, numbers of woody seedlings and ground cover of vascular plants relative to initial values were similar on cleared and uncleared treatments. Volume of plant biomass <2 m remained low on cleared subplots. Ungulates significantly reduced the re-establishment of woody seedlings ≥ 10 cm tall: only one seedling reached this height outside exclosures, compared with 29 seedlings inside. The number of seedlings <10 cm tall, expressed relative to numbers present pre-clearing, was not significantly affected by ungulates. The species composition of regenerating vegetation was more similar (Jaccard index) to pre-clearing understorey vegetation inside ungulate exclosures than outside. No consistent effect of rodents (primarily house mice) on seedling establishment or species composition was detected after 2 years, and rodent exclosures did not significantly affect survival of seedlings (Griselinia littoralis and Aristotelia serrata) planted as an index of rodent herbivory pressure. No significant differences in vegetation recovery were apparent between forest types. Rapid seedling recruitment in the absence of understorey vegetation and the presence of herbivores provided evidence that understorey vegetation competes with seedlings for light. Ungulate effects were consistent with other experiments that showed herbivores reduced the rate and altered the trajectory of vegetation regrowth after disturbance.     

Importance of animal and plant traits for fruit removal and seedling recruitment in a tropical forest

The traits of animals and plants influence their interaction networks, but the significance of species' traits for the resulting ecosystem functions is poorly understood. A crucial ecosystem function in the tropics is seed dispersal by animals. While the importance of species' traits for structuring plant–frugivore networks is supported by a number of studies, no study has so far identified the functional traits determining the subsequent processes of fruit removal and seedling recruitment. Here, we conducted a comprehensive field study on fruit removal by frugivorous birds and seedling recruitment along an elevational gradient in the Colombian Andes. We measured morphological traits of birds (body mass, bill width, Kipp's index) and plants (plant height, crop mass, fruit width and seed mass) which we expected to be related to fruit removal and seedling recruitment. We tested 1) which bird and plant traits influence fruit removal, and 2) whether network metrics at plant species level, functional identities of frugivores (community‐based mean trait values) and/or plant traits were the main determinants of seedling recruitment. We found that large‐bodied bird species contributed more to fruit removal than small‐bodied bird species and that small‐sized fruits were more frequently removed than large‐sized fruits. Small plant species and plants with heavy seeds recruited more seedlings than did large plants and plants with light seeds. Network metrics and functional identities of seed dispersers were unrelated to seedling recruitment. Our findings have two important implications. First, large birds are functionally more important than small birds in tropical seed‐removal networks. Second, the detected tradeoff between fruit size and seed mass in subsequent recruitment processes suggests that the adaptability of forest plant communities to a loss of large frugivores is limited by life‐history constraints. Hence, the protection of large‐bodied frugivores is of primary importance for the maintenance of diverse tropical plant communities.     

Consequences of habitat disturbance on seed fate of a Brazilian tropical dry forest tree Cavanillesia arborea (Malvaceae)

We compared seed fate (survival, mortality by rodent predators, desiccation and other causes) of the tree Cavanillesia arborea (Malvaceae) in preserved old‐growth tropical dry forests and in nearby abandoned pasturelands in Brazil. For this purpose, we performed an experiment where 15 seeds were placed in the surroundings of 15 parental individuals in each habitat. These 450 seeds were monitored over four months. At the end of the experiment, seed predation by rodents was higher in forest (56.5%) than in pasture (8.9%) areas, but seed desiccation showed the opposite pattern (8.9% vs. 80.4%). Mortality by desiccation was also faster in pasture than in forest areas, probably reducing their attractiveness to predators in these areas. None of the seeds placed in the pasture survived, whereas 26.2% of seeds became seedlings in forest areas. The absence of seedling recruitment of Cavanillesia arborea in pastures is likely a consequence of their incapacity to tolerate the harsh abiotic conditions in this habitat. Thus, forest conversion imposes a strong limitation to the long‐term population viability of this species. However, restoration and natural regeneration of abandoned pastures can recreate the forest structure and microclimatic conditions favourable to seed germination and seedling establishment. Remnant individuals of Cavanillesia arborea in agricultural landscapes may have a negligible contribution to current recruitment, but they can attract potential dispersers of pioneer species, with positive feedbacks to future recruitment during secondary succession.     

Large herbivores regulate the spatial recruitment of a hyperdominant Neotropical palm

Large mammalian herbivores play an important role in shaping the diversity of tropical forests by affecting the survival of seedlings and saplings beneath parent plants. The white‐lipped peccary (Tayassu pecari) accounts for the largest herbivore biomass that controls seed and seedling survival in Neotropical ecosystems. However, hunting and habitat loss has driven peccaries to local extinction for most of their original distribution, so it is likely that their absence will affect plant recruitment dynamics. We tested the effects of peccary local extinction on the density and spatial distribution of the hyperdominant palm Euterpe edulis by performing a fine‐scale characterization of its spatial recruitment in six forest sites in the Brazilian Atlantic forest. We compared the age structure and the spatial patterns of seedlings, saplings, and adults as well as the relationship between them. We found that while under the presence of peccaries there was a decrease in recruitment rates under adults, the local extinction of these large mammals led to a more clumped process of spatial recruitment. Despite such contrasting spatial patterns of recruitment dynamics, neither age structure nor the random spatial distribution of adults was affected by the presence or absence of peccaries, indicating that their early effects on these palm populations are mitigated as recruitment advances. Our findings highlight the role of large‐bodied forest‐dwelling herbivores in regulating the fine‐scale spatial recruitment of plants and advance our understanding on the effects of defaunation in tropical forests. Abstract in Portuguese is available with online material.     

Herbivores limit the population size of big‐leaf mahogany trees in an Amazonian forest

The Janzen–Connell hypothesis proposes that specialized herbivores maintain high numbers of tree species in tropical forests by restricting adult recruitment so that host populations remain at low densities. We tested this prediction for the large timber tree species, Swietenia macrophylla, whose seeds and seedlings are preyed upon by small mammals and a host‐specific moth caterpillar Steniscadia poliophaea, respectively. At a primary forest site, experimental seed additions to gaps – canopy‐disturbed areas that enhance seedling growth into saplings – over three years revealed lower survival and seedling recruitment closer to conspecific trees and in higher basal area neighborhoods, as well as reduced subsequent seedling survival and height growth. When we included these Janzen–Connell effects in a spatially explicit individual‐based population model, the caterpillar's impact was critical to limiting Swietenia's adult tree density, with a > 10‐fold reduction estimated at 300 years. Our research demonstrates the crucial but oft‐ignored linkage between Janzen–Connell effects on offspring and population‐level consequences for a long‐lived, potentially dominant tree species.     

Combined effects of seed and soil quality drive seedling performance of a late‐successional canopy tree in a tropical forest

Habitat loss and fragmentation affect the structure and functioning of forested ecosystems worldwide, yet we lack an understanding of how species respond to environmental changes. Here, we examined reproductive success and seedling performance of Poulsenia armata (Moraceae) in continuous and fragmented forests of Los Tuxtlas, southern Mexico. We further investigated how maternal habitat and soil conditions manifested in the seedling stage. We determined seed quality and seedling performance by combining isotopic analyses in seed quality with field observations of P. armata fruit production and a common‐‐garden experiment. Soil conditions in forest fragments negatively impacted P. armata reproductive success. Trees of P. armata in forest fragments were smaller in size and produced fewer fruits and smaller seeds with lower quality compared with trees from the continuous forest. The combined effects of maternal habitat and soil conditions determined seedling survival and growth of this tropical tree. Notably, seedlings had restricted plasticity for biomass allocation to roots, limiting the capacity of fragmented populations to compensate for the initial low N content in seeds. Trees in forest fragments at Los Tuxtlas produced offspring competitively inferior and potentially less resilient than counterparts in continuous forest, jeopardizing future persistence of this late‐successional tree species.     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.